Contributions
'of the American Entomological Institute
Volume 24, Number 3,1988
PISON IN THE NEW WORLD: A REVISION (H...
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Contributions
'of the American Entomological Institute
Volume 24, Number 3,1988
PISON IN THE NEW WORLD: A REVISION (HYMENOPTERA: SPHECIDAE: TRYPOXYLlNI)
by
Arnold S. Menke
Pison in the New World: a revision (Hymenoptera: Sphecidae: Trypoxylini) By Arnold S. Menke Systematic Entomology Laboratory Agricultural Research Service, USDA c/o U. S. National Museum, Washington DC 20560 ABSTRACT The genus Pison Jurine in the Western Hemisphere is revised. Identification keys are provided for the forty four recognized species, all but one of which are restricted to the Neotropical Region. All species are described, diagnosed, and their geographic ranges outlined. Illustrations accompany the key and descriptions. Twenty nine new species are described: abathes (Ecuador, Bolivia, Guyana), abothrum (Colombia, Brasil), arachniraptor (Panama to Bolivia, Brasil), aranevorax (Colombia, Ecuador, Peru, e Brasil), brasilium (Brasil). chrysops (Costa Rica to Argentina), cooperi (Costa Rica to n Bolivia. c Brasil, Dominica), delicatum (South America), dementia (se Brasil), doggonum (Mexico), eu (Mexico to Peru & Suriname), euryops (Colombia, Brasil, Argentina), erebus (Colombia), eyvae (Colombia, Ecuador, Bolivia), fritzi (Ecuador, Brasil, Argentina), gnythos (Colombia, Ecuador, Guyana, Trinidad), larsoni (Ecuador, Peru, Bolivia), lillo (Argentina), longicorne (Mexico to Argentina), martini (Ecuador). nosferatu (Venezuela), oaxaca (Mexico), pentafasciatum (s Brasil), phthinylla (Ecuador), sphaerophallus (Colombia, Ecuador, Peru, Guyana, Suriname, n Brasil), styphopteron (Colombia, Peru), ~ (Peru), vincenti (Ecuador, Guyana), and wasbaueri (Argentina). One species is synonymized: flavolimbatum Turner, 1917 = cressoni Rohwer, 1911. Pison laeve Smith, known only from its type specimen and sometimes treated as a North American species, is interpreted as an Australasian taxon and a diagnosis based on the type is presented. Twelve species groups are established for the New World fauna, but the use of subgenera is abandoned. Three generic names that have been treated as subgenera in the past, Pisonoides Smith, Krombeiniellum Richards, and Entomopison Menke are placed in synonymy with Pison. Some characters that are important from a phylogenetic standpoint in Pison and related genera are analyzed to determine polarity. Incorporation of the Crabroninae with the Larrinae is discussed. The status of Pisonopsis as a genus is reviewed, a new generic character described, and a key to its five species is presented. Two new species of Pison are described from New Guinea that are important for an understanding of generic limits: woji and pistillum. CONTENTS Introduction . . . . . . . . . . . . Acknowledgements
. .
Methods and Techniques
2
Distinctions between Pison and Pisonopsis
3
Genus Pison Jurine
5
Subgenera . .
5
ii
Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988 Species groups
. .
7
Species characters
8
Character analysis
13
Biology
18
. . . . .
Key to species of New World Pison
19
Con forme group . . . conforme Smith doggonum Menke eyvae Menke larsoni Menke
24 25 26 27
Fritzi group . . . . fritzi Menke . ;:;QSf8ratu Menke
30 31 32
Eremnon group eremnon Menke
33 33
Delicatum group. . . delicatum Menke
35 35
Agile group . . . agile (Smith)
37 38
Stangei group . . plaumanni Menke stangei Menke duckei Menke abathes Menke Krombeini group . . krombeini Menke neotropicum Menke Euryops group . . . eu Menke . . euryops Menke lillo Menke StYPhopteron Menke Cressoni group cressoni Rohwer chrysops Menke erebus Menke . P8iit1ifasciatum Menke martini Menke . . maculipenne Smith . brasilium Menke . . arachniraptor Menke cameronii Kohl dementia Menke . abothrum Menke . aranevorax Menke phthinylla Menke
29
39
40 41 42 43
44 45 46 46
47 48
49 50 51 53 55 57 58 58 59 61 62 64 66 67
68 70
Menke: Pison in the New World
iii
Chilense group. . . . chilense Spinola ~Menke. .
71 71 73
Convexifrons group. . convexifrons Taschenberg . wasbaueri Menke. . . cooperi Menke . . longicorne Menke.
74 75 76 77 78
Pilosum group . . . . . pilosum Smith . . aureofaciale Strand. vincenti Menke. oaxaca Menke . . . gnythos Menke . . . sphaerophallus Menke
80 81 84 84 85 86 88
. . . . . . . . . . . . . . . . . . . . . . ..
Appendix. . . . . . .
90
Pison laeve Smith
90
New species of Pison from New Guinea pistillum Menke . . . woji Menke. . . . . .
90 91 92
Key to species of Pisonopsis
93
Literature cited
94
Illustrations . .
97
INTRODUCTION The spider predator genus Pison Jurine is cosmopolitan, but in the Western Hemisphere it is essentially a neotropical taxon. All but one of the 44 species treated here are restricted to that region. The only nearctic species, agile (Smith), is not native; it was introduced into northeastern North America from the Orient during World War II. Another species attributed to the North American fauna, laeve Smith, appears to belong in the Indoaustralian area based on my examination of its type (see Appendix, p. 90). This study has been based on 1580 specimens of Pison from the Western Hemisphere, of which 484 were chilense Spinola, a commonly collected endemic in Chile and Argentina. When I began my review of the genus, only 15 neotropical species were known, but I now recognize 43 species in the region, 29 of them new. This swells the world total for Pison to somewhere around 200 described species making it one of the larger genera of Sphecidae. More neotropical species await detection, perhaps as many as 20. and of course the large Australian fauna has yet to be subjected to a modern review. Worldwide, the genus may eventually contain nearly 300 species. This revision has to be considered as only a beginning framework for several reasons. Some species are known only from one sex or even one specimen. I have studied a number of specimens that represent additional undescribed species, but I have not attempted to describe them because they are singletons and are in a species group (the cressoni group) whose species are often difficult to distinguish even when a lot of material is available. Furthermore, several species treated here may ultimately prove to be complexes. Finally, in a few instances I have been able to separate males, but their corresponding females seem identical. Clearly much more collecting is needed before a definitive revision can be achieved. Such a revision will also require rearing of nest material to positively associate sexes. I have not attempted a comprehensive phylogenetic analysis of the New World fauna of Pison. Such an analysis would have to be considered preliminary because the very large Australian fauna (about 50 named species of Pison), which may have some Gondwanian links with the South American fauna, is poorly known, and the rest of the world species of the genus are not well documented morphologically. Lack of a comprehensive world collection of Pison, as well as time constraints imposed on my study, have precluded an in-depth world survey of the genus. I have analyzed a number of characters, however, that are phylogenetically and morphologically important in the genus, in the Trypoxylini, as well as in the Larrinae. I have concluded that the Crabroninae should be incorporated with the Larrinae. In the Appendix at the end of this paper I describe two new species of Pison from New Guinea that are important from the standpoint of understanding the morphological diversity in the genus. A key to the five species of the New World genus Pisonopsis Fox, a taxon separated with difficulty from Pison, is also found in the Appendix. ACKNOWLEDGMENTS During the writing phase of this project I sought and received advice, OplnlOnS and counsel from a number of people: Wojciech Pulawski, J'ames Carpenter, E. Eric Grissell, Maria Alma Solis, David Wahl, Richard Bohart and Karl Krombein. Their thoughts and help are appreciated. Drafts of the manuscript were critically reviewed by Wojciech Pulawski, James Carpenter, and Ray Gagne and their comments enabled me to considerably improve the ms. Duane Hope demonstrated how to mount genitalia on gold wires with polyvinyl acetate (PV A) for SEM photomicrography. Brian Kahn, Susann Braden, and Walter Brown came to my assistance whenever I ran into problems on the scanning electron microscope. Leslie Brothers provided a provincial map of Ecuador. Bonnie Farmer corrected bird names cited in biology sections. Vera Lee assisted in preparing camera ready copy. To all these people I extend my sincere thanks.
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Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988
Colin Vardy of the British Museum (Natural History), Max Fischer of the Naturhistorisches Museum, Vienna, the late Eberhard Konigsmann of the Humboldt Museum, Berlin, and J. O. Husing of the Martin Luther UniversiUit, Halle, generously lent type material. Other research material was borrowed from the following collections. Abbreviations used in the species treatments are listed first. AEI - American Entomological Institute, Gainesville (Henry Townes). AMNH - American Museum of Natural History, New York (Marjorie Favreau). BERLIN - Zoologischen Museum der Humboldt Universitat, Berlin (Eberhard Konigsmann, F. Koch). BMNH - British Museum (Natural History), London (Colin Vardy, Michael Day). BPBM - Bernice P. Bishop Museum, Honolulu (Gordon Nishida). CAS - California Academy of Sciences, San Francisco (Wojciech Pulawski). CMP - Carnegie Museum, Pittsburg (George Wallace). CNC - Canadian National Collection, Ottawa (Lubomir Masner). COOPER - Martin Cooper Collection, Lyme Regis, Dorset, England CSDA - California State Department of Agriculture, Sacramento (Marius Wasbauer). CU - Cornell University, Ithaca (George C. Eickwort). DIAS - Braulio F. de Souza Dias Collection, Fundacao IBGE, Brasilia. FRITZ - Manfredo Fritz Collection, Salta, Argentina. FSDA - Florida State Department of Agriculture, Gainesville (Lionel Stange). GEMBLOUX - Faculte des Sciences agronomiques de l'Etat, Gembloux (Jean Leclercq). GENEVA - Museum d'Histoire Naturelle, Geneva (Claude Besuchet). HALLE - Martin Luther Universitat, Halle, East Germany (J. o. Husing). HEH - Henry Hespenheide Collection, Los Angeles. IBGE - Fundacao IBGE, Brasilia (Braulio de Souza Dias). KU - University of Kansas, Lawrence (George Byers, Robert Brooks, John Wenzel). LECLERCQ - Jean Leclercq Collection, Liege, Belgium (Jean Leclercq). LEIDEN - Rijksmuseum van Natuurlijke Historie, Leiden (Kees van Achterberg). LILLO - Instituto Miguel Lillo, Tucuman (Abraham Willink, Lionel Stange). MCZ - Museum of Comparative Zoology, Cambridge, Mass. (James Carpenter). MLSU - Zoological Museum of the Moscow Lomonosov State University, Moscow (Alexander V. Antropov). OSU - Oregon State University, Corvallis (George Ferguson). PARIS - Museum National d'Histoire Naturelle, Paris (Simone Kelner-Pillault). PMA - Provincial Museum of Alberta, Edmonton (Albert Finnamore). RAW - Anthony Raw Collection, Brasilia. UCD - Richard M. Bohart Museum, University of California, Davis (Robert Schuster). UCM - Universidad Central de Venezuela, Maracay (F. Fernandez-Yepez). USNM - National Museum of Natural History, Washington DC. (Arnold Menke). USU - Bee Biology and Systematics Lab., USDA, Utah State University, Logan (Terry Griswold). VIENNA - Naturhistorisches Museum, Vienna. (Max Fischer) ZMC - Universitetets Zoologiske Museum, Copenhagen (Ole Lomholdt). METHODS AND TECHNIQUES I have not provided complete label data for all species. For common, described taxa I have given only the country, province, and locality followed by the abbreviation of the owner of the material in parentheses. The number and sex of specimens from anyone locality is usually given only for new species; males are noted however, for species in which that sex is poorly represented. Collectors names are cited only for material of new species. Some of my new species may prove to represent complexes of two or more species. In these cases I have not always included all material studied in the para type series. Instead atypical specimens are listed as meta types, i.e., non-type material.
Menke: Pison in the New World
3
Extensive use of scanning electron photomicrographs has been made to illustrate various body parts. Genitalia and a few other structures were coated with gold palladium, but most material was photographed uncoated, particularly clypei, mandibles, and propodei. A Cambridge Stereoscan 100 set at 10 KV was used for coated material. Uncoated material was scanned at 2 KV on the same machine using a low KV anode, or occasionally on a Hitachi S-570. Uncoated specimens were held in a special stub similar to that described by Stork and Claugher (1987). This stub (fig. I) permits leaving specimens on their pins or points during SEM work, and allows photographs of holotypes and unique material to be made without damage. Uncoated material was often soaked in an undiluted solution of the commercial cleaning agent, 409, and then run through water, alcohol, and chloroform. The last causes setae to remain erect after drying. Sometimes cleaning in chloroform was enough. Debris was removed from the body by using a small piece of Scotch Magic Transparent Tape stuck to the end of an insect pin. Dust and other small particles adhere to the tape. Male genitalia and sternum VIII were glued to the end of fine gold wire with polyvinyl acetate (PVA) dissolved in a mixture of acetone and 99% isopropyl alcohol. The wires were then glued to standard SEM stubs with Elmer's Glue-All (fig. 2). These structures were not cleared, but were often cleaned in an undiluted solution of 409 cleaning agent. The PVA was usually nearly dry on the wire by the time it was applied to a specimen, so to facilitate adhesion of these small objects they were soaked in 99% isopropyl alcohol immediately before touching the wire to them. A Wild M-400 photographic microscope was used to take pictures of wings and some propodei.
Figures 1-2. SEM mounting techniques. I, stub drilled to accept insect pin to permit photography of uncoated type specimens and other unique material. 2, male genitalia and sternum VIII mounted on gold wires that are glued to standard stub. DISTINCTIONS BETWEEN PISON AND PISONOPSIS These two genera are separated by rather tenuous differences (see Bohart and Menke, 1976:330). Pisonopsis has a notched mandible, but so do some Pison. However, Wojciech Pulawski (in !itt.) has discovered another mandibular feature that ultimately may be the only character that will permit maintaining E1sonopsis as a discrete genus. In Pison the flattened lateral face of the mandibular base is margined posteriorly (or below depending on your viewpoint) by a rather sharp edge (condylar ridge of Michener and Fraser, 1978) that extends from the lower condyle to the notch (figs. 4-6), or to the midpoint of the much more common notchless mandible (fig. 3). This condition occurs in all other trypoxylines and Larrinae that I have examined
4
Contrib. Amer. Ent. Inst.. vol. 24. no. 3. 1988
except Pisonopsis. In Pisonopsis the lateral face of the mandible has a rounded ridge that parallels the condylar ridge of the mandible delimiting a groove (figs. 7-8). This secondary structure is best developed in clypeata Ashmead. areolata (Spinola). and australis Fritz. and it appears to be an autapomorphy for Pisonopsis. Clearly the mandible in the Sphecidae needs further study since it may offer defining features not heretofore appreciated. The graduli found on abdominal sterna III-IV of Pisonopsis do not occur in Pison. but a practical problem of using them as a generic character is that they are often barely visible due to telescoping of the abdominal segments. and they are especially
Figures 3-8. Left mandible in lateral view. 3-6. Pison species (arrow indicates condylar ridge); 3 is arachniraptor. 4 is species from New Guinea. 5 is convexifrons. 6 is gnythos. 7-8. Pisonopsis species (arrow indicates secondary ridge); 7 is birkmanni. 8 is areolata.
Menke: Pison in the New World
5
weak in areolata from South America. The marginal cell of Pisonopsis is rounded apically, rather than acuminate as in most Pison. But the cell is rounded also in some Pison (eremnon for example). The basal flagellomeres of the male antenna in Pisonopsis are asymmetrically swollen in all species. The asymmetry of the first or first and second flagellomeres is rotated 90 to 180 0 in relation to flagellomeres II or III-V or VI. Some species of Pison have asymmetrical flagellomeres also, and it is not clear if the male antennal structure in Pisonopsis can be used as a generic character. This needs more study. Since no key is available for the five species of Pisonopsis, I have included one in the Appendix (p. 93). This key was partly developed by R. M. Bohart but has been revised by me and expanded to include the South America species. GENUS PISON JURINE Pison Jurine (in Spinola), 1808:255. Type species: Pison jurini Spinola, 1808, monotypic. Pisonoides F. Smith, 1858: 104. Type species: Pison obliteratum F. Smith, 1858, monotypic. NEW SYNONYMY. Krombeiniellum Richards, 1962: 118. New name for Paraceramius Radoszkowski, 1887: 432. Type species: Paraceramius koreensis Radoszkowski, 1887, automatic. NEW SYNONYMY. Entomopison Menke. 1968a:1. Type species: Pison pilosum Smith, 1873, original designation. NEW SYNONYMY. The generic description in Bohart and Menke (1976:332) adequately characterizes Pison, but one character was overlooked. A nearly universal feature of the genus is a pit or transversely elongate depression anteromedially on the pronotum (figs. 16, 95). This is usually obscured by the back of the head, but it is an important apomurphic trend in Pison. I examined about 50 Old World species of Pison and only four were found to lack a pronotal depression. These were obliteratum Smith, xanthopus (Brulle), multistrigatum Turner, and kohlii Bingham. In the New World a pit is absent in the euryops and krombeini groups and a few species in the cressoni group. Trypoxylon and Pisoxylon apparently always lack these depressions. Species of Pisonopsis and Aulacophilus have either a pit or transverse depression. I have not seen material of Aulacophilinus. The discovery of two unusual species in New Guinea (see Appendix, p. 90) complicates the traditional generic concept. These two taxa have an unusually elongate clavi form first abdominal segment (figs. 347-350), and at first glance look like members of Trypoxylon. But both have three submarginal cells and two recurrent veins in the forewing (figs. 345-346), and an anterodorsal pronotal pit. Unlike nearly all Pison species, these two wasps have enlarged eye facets beneath the eye notch (figs. 341-344), a universal feature of Pisoxylon and most members of Trypoxylon s. s. Interestingly the facets are fairly uniform in size in most (all?) species of the subgenus Trypargilum of Trypoxylon. The antennal sockets are contiguous with the clypeus in one of the species of Pison from New Guinea, and slightly above the frontoclypeal suture in the other. The sockets are far above the clypeus in Pisoxylon and Trypoxylon except for some groups within the subgenus Trypargilum. Pison lobiferum Arnold from Madagascar (Arnold, 1945) has three submarginal cells and a clavate first abdominal segment but its wing venation is quite different, and it apparently lacks a dorsolateral propodeal ridge. Thus a close relationship to the two New Guinea wasps seems unlikely. SUBGENERA Four subgenera were recognized by Bohart and Menke (1976): Pison s.s., Pisonoides, Krombeiniellum and Entomopison. I now believe that subgenera are unwarranted as discussed below.
6
Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988
Pisonoides was proposed (Smith, 1858) for Pison obliteratum, an Indonesian species with two submarginal cells and a clavate first abdominal segment, but Turner (1916) used it for all species of Pison with two submarginal cells although he was aware that it was not a monophyletic sUbgenus. Two-celled wings have arisen independently a number of times in unrelated species or groups of Pison, and I (Menke, 1968a) restricted Pisonoides to three species with a clavate abdomen and two submarginal cells (obliteratum, difficile Turner and icarioides Turner). Lomholdt (1985: 15) treated Pisonoides as a genus and further refined its definition by including only species (unspecified) with a "constriction" between abdominal segments I and II. Lomholdt (1985:4) listed Pisonoides exclusum Turner as the species he studied but Turner (1916) described its first abdominal segment as not elongate. I have studied obliteratum and difficile, and based on their overall morphology they have convergently developed two-celled wings and an elongate gaster and are not closely related. Thus there is no reason to maintain Pisonoides and I am synonymizing it under Pison. Pison difficile is of interest in that it is one of the very few species of the genus with a noncarinate hindcoxa (see p. 17). Krombeiniellum has been used for species in which the eyes are densely microsetose (fig. 9), and was originally used for taxa with only two submarginal cells. I (Menke, 1968b) expanded the subgeneric concept to include species with setose eyes and three submarginal cells. My review of the neotropical fauna of Pison has shown that the three-celled species are not closely allied to the two-celled forms and that there are two disparate groups of species with two-celled forewings (agile and stangei groups). Furthermore, Pison delicatum, a new species with asetose eyes and three submarginal cells, shares several features with the agile, stangei and kr9mbeini groups: a short, clavate antenna; nonspiny legs (fig. 72); a velvety covering of short body setae; and thick tarsal claws (fig. 70). If Krombeiniellum is to be recognized as a subgenus, it must be redefined on the basis of these latter features, some of which are apomorphies. I am not certain, however, that this would result in a monophyletic taxon and am simply synonymizing the subgenus with Pison. Two neotropical species groups of Pison, the convexifrons and pilosum groups, are unusual in that the mandible has a notch on the posterior surface (figs. 5-6). I (Menke, 1968a) established the subgenus Entomopison for these species. This taxon has two apomorphies: the notched mandible and the polished, asetose "bearing" surface on the anterior margin of the pronotum (figs. 288, 295). The mandibular notch was thought to be unique to these New World species until Wojciech Pulawski returned from New Guinea in 1987 with a Pison (CAS), probably undescribed, that has a small notch. Actually the notch in this wasp is more like a "step" (fig. 4) that is delimited basad by a tooth-like angle. A similar but smaller step is present on the mandible of the Australasian and western Pacific island inhabitant. Pison ignavum Turner. Neither of these species have a broad pronotal bearing surface. In the two New World species groups this pronotal structure takes on two very different forms: a mediodorsal expansion of the polished rim that is not free (fig. 288, convexifrons group), and a broad lamella that is free from the rest of the notum (fig. 295, pilosum group). The two types represent convergent specializations in my opinion. Similar lamelliform structures are found in two other neotropical groups of Pison, the fritzi and krombeini groups, but in these the surface is setose, at least in part, and not always highly polished (fig. 45). Based on the foregoing, the convexifrons and pilosum groups probably represent two unrelated monophyletic lineages and I have decided to synonymize Entomopison under Pison. Finally, Antropov and Pulawski (in prep.) are describing a fossil species of Pison from Baltic amber that has a notched mandible similar to that found in the convexifrons and pilosum groups. I have examined the unique specimen, and it appears to have an elongate anterodorsal pronotal pit that is bordered anterad by an overhanging lamelliform carina. It is impossible to determine if there is a polished bearing surface as in the convexifrons group, but there is no free lamella like that found in the pilosum group. The labrum appears quadrangular and is not emarginate, and the occipital carina is incomplete ventrad, ending well before the hypostomal carina. In these respects the fossil is like the convexifrons group. The pitted
Menke: Pison in the New World
7
episternal sulcus is straight. ending ventrad before curving forward. The metapleural flange looks like it is somewhat lamelliform. and almost as broad as a hindocellus. Hindtarsomeres II-IV each have a plantula. The propodeal dorsum has ridges that extend from the base in an oblique fashion. and a median. longitudinal carina . The fossil differs from present day species of the convexifrons and pilosum groups in at least two ways: the propodeal side is not delimited above by a crenulate ridge or line of foveae. and the second recurrent vein terminates on the third submarginal cell. Clearly the fossil represents a third lineage with a notched mandible. SPECIES GROUPS The 44 New World species segregate into 12 groups. all but one of which (the conforme group) are defined by clear apomorphic features. Most of these groups appear to be endemic to the New World. but the agile group is an Old World assemblage with an introduced species in North America. The chilense group may have relatives in Australia. Some of the characters useful in defining groups include: form of the mandible (position of teeth. presence of notch); shape and details of clypeal lobe; shape of labrum; form of the occipital carina; whether or not the frons is strongly swollen; proportions of head (eye length vs. distance between eye notches); presence or absence of dense setae on eye; presence or absence of an anterodorsal lamella or boss on the pronotum; presence or absence of a crenulate ridge at the top of the propodeal side; sculpture of propleuron; form of the episternal sulcus; wing venation; and form of male sternum VIII and genitalia. The presence on the eye of short . dense setae is an apomorphic trait common in several groups (fig. 9). but some species of Pison have a few short setae widely scattered over the eye (cressoni group for example). In general a naked eye is the common condition.
Figure 9. Left eye of Pison plaumanni showing eye setation.
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Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988
The form of male sternum VIII is fairly uniform within a species group. It may be simply truncate or rounded apically (fig. 17), the plesiomorphic state, or the apex may be emarginate (fig. 238). The emargination is often bounded laterally by a blunt prong, or pseudosting (fig. 254). The makeup of the male genitalia varies among the 12 species groups, some being robust, some laterally compressed, and some more or less dorsoventrally compressed. The gonostyle is a simple structure in the conforme, fritzi, eremnon, delicatum, agile, and euryops groups. In the pilosum group the gonostyle is often a simplified, large plate-like affair. The gonostyle has a ventral lobe in the krombeini, cressoni, and convexifrons groups, and there are two ventral lobes in the chilense group. I regard the presence of one or two gonostylar lobes as an apomorphy. The volsellar lobes are very large in the conforme, fritzi, eremnon, krombeini, and cressoni groups, but are quite small in the agile, stangei, convexifrons and pilosum groups. In the eremnon group, the volsellar lobes are represented by two long, narrow structures. Descriptions and discussions of apomorphies are provided for most of the species groups. Descriptions have not been given for the two monotypic groups (eremnon and delicatum), but their species descriptions and discussions are adequate. SPECIES CHARACTERS Described here are features used in species descriptions and group diagnoses that need clarification or merit discussion. Morphological terms are from Bohart and Menke (1976) except that for surface sculpture I have used the terminology developed by Harris (1979). I have used the terms "carina" and "ridge" more or less interchangeably. A lamelliform carina is one that is elevated into a thin blade-like structure. Structures that have not been studied carefully for use in species discrimination are mouthparts, proportions and details of leg segments especially the tarsi, and details of wing venation such as proportions of cells and cell veinlets and the form of individual veins and cells. Apparent differences in the form of the propodeum (length, angle between dorsum and hindface, etc.) have been noted in some species but these are difficult to quantify and have usually been ignored. Labrum: Sometimes the labrum is short, broad and is described as "transverse". Most often the labrum is "quadrangular", i.e., more or less truncate apically with perpendicular sides. It is deeply emarginate in the euryops and pilosum groups with the result that usually only two finger-like lobes are visible beneath the clypeal margin. Clypeus: The clypeus is one of the more important structures for species discrimination. The free edge often has an impunctate, asetose rim or "lip", and usually projects as a median lobe whose shape varies from triangular to truncate or rounded. The lobe may bear several "teeth". When viewed from below, the free edge of the clypeus is often clearly "thickened" or "double-edged". In the latter instance the thickening is margined behind by a carina. Often the thickest part is the section between the median lobe and the lateral end point of the clypeal margin. The clypeal surface is often covered by metallic silver setae that are appressed and obscure the surface sculpture. Mixed with these short setae are longer setae that are directed downward, and in some cases they are quite bristly and form a "brush" that surpasses the edge of the clypeus (fig. 6). [rons: The common appearance is a gently convex surface, but in the convexifrons group the frons is strongly swollen toward the antennal sockets. In the pilosum group some species have a small, impunctate polished depression or "dimple" on the midline above the sockets. This feature is large in gnythos, new species. Antenna: The antenna varies from moniliform with elongate flagellomeres (two or m~re times as long as wide) to clavate where the flagellomeres are progressively
Menke: Pison in the New World
9
shorter and broader toward the apex. Measurements of length and width of flagellomeres were made with an ocular micrometer at a magnification of SOX . Width measurements were always taken at the apex of an article, and the microsetae on the surface were not included. The male antenna sometimes has tyli or other modifications on some of the f1agellomeres. When present, tyli occur on the surface that is on the inside of the antennal curl, and great care must be used in examining the flagellum for the presence of these and other specialized structures. Good light and high magnification (SOX) are essential, and often the antenna has to be relaxed and uncurled in order to see the surface bearing these structures. A "simple" antenna is one in which the flagellum has no visible modifications. Head measurements: The "upper interocular distance", or UID, is the shortest distance between the eyes at the vertex - it is always measured behind the ocellar triangle. The "lower interocular distance", or LID, is the shortest distance between the eyes adjacent to the clypeus (or near level of antennal sockets in some extralimital species). Eye length is measured vertically and is compared with the distance between the eye notches on the inner orbits (fig. 10). The "ocellocular distance", or ODD, is the measurement made between the inner orbit of the eye and the hindocellus. This is compared to the "hindocellus diameter" , or HOD. All of these measurements were made with an ocular micrometer at a magnification of SOX to maximize accuracy.
Figure 10. Face of Pison eu, female, showing how to measure eye length and distance between eye notches. -
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Contrib. Amer. Ent. lnst., vol. 24, no. 3, 1988
Occipital carina: This is usually "incomplete", i.e, it does not form a complete circle. The lower ends of the carina either terminate just before the midline of the head, or just before meeting the hypostomal carina, or sometimes actually meet the hypostomal carina. Pronotal pit and lamella: Because the anterior portion of the pronotum is usually hidden under the rear of the head, its structure has not been studied, at least in the Sphecidae. But in Pison I discovered two types of features that merit the attention of anyone working on the family. One character is the development in some taxa of the narrow polished anterior rim into either a broad boss or a broad, free lamella. The second is the presence of a round pit or transversely lengthened depression behind the anterior margin of the pronotum. Both features can only be viewed adequately when the head is tilted forward. The normally narrow polished rim of the pronotum is occasionally broadened dorsally into a polished boss (convexifrons group, fig. 288) that is suggestive of a bearing surface for the back of the head (this may not be its true function). The more usual modification however, is the extension of the polished rim upward and backward as a broad, free lamella. In the pilosum group it extends laterad to about the level of the pronotal lobe (figs. 294-295), and the area directly beneath it is smooth and polished. Sometimes this lamella is present laterally but narrows toward midline where it is paralleled posteriorly by a lamelliform, setose carina that borders the anterodorsal pro notal depression (fig. 45, fritzi n. sp.). The lamella may be asetose and polished (pilosum and fritzi groups) or minutely setose (krombeini group). The "anterodorsal pit of the pronotum" varies from a small round pit whose diameter is less than that of a hindocellus (fig. 95), to a "transversely elongate depression" (fig. 67) that mayor may not be bordered anteriorly by a sharp edge or carina, the latter sometimes lamelliform, dull, and setose (delicatum, fritzi, n. spp.). When lamelliform this carina can be confused with the anterodorsal lamella described above, but the latter is usually polished and often asetose. When the pit is transformed into an elongate depression its surface is impunctate and shiny. When I refer to its "length" in descriptions and discussions, I mean its transverse length, Le., perpendicular to the longitudinal axis of the body. Pronotal collar: This is a fairly simple structure in most species, but it may be thick or thin and comparison to the width of the metanotum is usually made. Occasionally the "humerus", or lateral corner of the collar, is angUlar, the angle being part of a transversely oriented ridge (fig. 99). In some species the collar has a median prominence (fig. 288), or the anterior face may be flattened at the middle. Scutal flange: The lateral edge of the scutum adjacent to the tegula is reflexed upward forming a flange of variable width. Lateral ridge of propodeum: Some groups of Pison have a ridge that extends in a gradual arc from the spiracle to the propodeal socket (fig. 27). Since it is situated on the interface between the propodeal dorsum and side, it delimits the top of the latter. Usually associated with the dorsal side of this ridge are cross-carinae of variable length that are perpendicular to it. This usually makes the ridge appear crenulate. Sometimes the cross-carinae are reduced to very short teeth along the dorsal side of the ridge. In some species, the ridge is replaced by a row of large foveae as in the pilosum group. In a few species the ridge (larsoni n. sp.) or row of foveae (pilosum Smith) vary in development from present to non-existant. Forewing: The marginal cell is typically acuminate distally. This is enhanced by the slight extension of vein RI beyond the end of the cell. In eremnon Menke the RS vein curves forward at the cell apex where it meets RI perpendicularly (fig. 53). A somewhat intermediate condition occurs in chilense (fig. 253). Although three submarginal cells is the norm in Pison, with the second petiolate, two-celled wings are characteristic of the agile and stangei groups (fig. 12), and are also found in a few species of other groups (fig. II). This reduction in submarginal cells occurs two
Menke: Pison in the New World
II
ways. Loss of the outer vein let (lr-m) of the second submarginal cell results in the two-celled condition seen in some species of the cressoni and euryops groups. In these the remaining veinlet is usually angulate (fig. II). Diminution (figs. 87-88) and eventual loss of the second submarginal cell is another way of achieving a two-celled wing (agile and ~tangei groups). Here the resulting veinlet between the remaining submarginal cells is more or less straight (fig. 12). The shape of submarginal cell III varies at the group level. The inner and outer veinlets (Ir-m, 2r-m) may meet on the marginal cell or be well separated there. In rare instances the two vein lets join before reaching the marginal cell resulting in a petiolate third submarginal cell. The termination point of the recurrent veins (lm-cu and 2m-cu) varies considerably in Neotropical Pison, sometimes even within a species. The point of divergence of the forewing media in relation to crossvein cu-a is useful in some species. Typically the media diverges after cu-a, but occasionally it is before cu-a as in abothrum and brasilium.
®
® Figures 11-12. Right forewing. Pison plaumanni Menke.
11, Pison styphopteron n. sp. (holotype).
12,
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Legs: The hindcoxa has one or two longitudinal carinae dorsally. The inner one is always present and sometimes it is partially lamelliform. Apparent differences between species in the degree and shape of the lamelliform part of this carina seem unreliable, however. An outer carina is present in the convexifrons and pilosum groups, but in most Pison it is usually present only on the apical one half, or less. It is absent in a few species. The male foretrochanter is usually simple, but it has a sharp spine in some species of the pilosum group (figs. 292-293). A plantula occurs at the ventral apex of tarsomeres II-IV in some species (figs. 13-14, fritzi and pilosum groups, chilense, eremnon), but usually only tarsomere IV has one, and they appear to be absent in the krombeini and stangei groups and in a few other species. Usually the legs have some conspicuous spines at the apex of the tarsomeres, but in the krombeini, agile, stangei and delicatum groups the legs are essentially unarmed (fig. 72). The claw is unusual in these latter forms also. Instead of gradually tapering to the apex, the claw is of fairly uniform width until just before the apex (fig. 70). The claws in males of the pilosum group are asymmetrical on the foreleg, and sometimes on the middle leg. In this group the anterior claw is deformed and lamelliform (fig. 14).
Figures 13-14. Details of male foretarsus of Pison pilosum. 13, ventral view of tarsomeres III-IV showing plantulae. 14, ventral view of tarsomeres III-V showing deformed claw.
Menke: Pison in the New World
13
Gaster: This is the definitive abdomen and the segments are numbered I-VI or VII depending on sex. Segment I is somewhat elongate in a few species and I have expressed this by measuring the length of tergum I along its midlength in dorsal view, and comparing that to the apical width. This is somewhat imprecise because the midlength can vary depending on how the segment is oriented. The first two or three terga are sometimes rather strongly convex with result that in lateral profile there is a strong constriction between terga I and II (pilosum group, fig. 301). In a few species of the cressoni group, especially in the female, tergum II has a median hump (figs. 162, 251). More commonly tergum II may have a pair of low swellings, one on each side of the midline, as in the pilosum group, eremnon, and others. The apical margin of tergum I is sometimes double-edged (euryops and pilosum groups, fig. 335), or it may be depressed subapically forming an apical band (eremnon), or the edge may be reflexed as in the krombeini group. Some species have tergal fasciae of setae (chilense group for example) or yellow bands (cressoni group). Male tergum VII is generally simple, but it is laterally compressed in some species of the convexifrons group, truncate apically in the chilense group, and ends in two rounded, reflexed lobes in eremnon. Tergum VI in eremnon is deeply V-notched apically (fig. 56). Although the sterna are sometimes modified in the females (slight median hump on sternum II in conforme, for example), specializations on the venter occur mainly in males and are presumed to have a role in copulation. The male of eremnon, which has the most sternal embellishments among New World Pison, has transverse flanges on sterna II-Ill, setal depressions on IV-VI, and sterna VI-VII are emarginate apically (figs. 58-59). The male of delicatum has decumbent mats of setae on sterna Ill-V (fig. 68). Sternum VII sometimes has depressions of various shapes as in the pilosum group (figs. 304-306). Sternum VlII may be truncate apically (fig. 108), rounded apically (fig. 18), emarginate apically (fig. 69), or end in two strong prongs (fig. 254). Male genitalia: These structures usually offer good specific differences in New World Pison and I have used scanning electron photomicrographs to illustrate them. The ventral side of the genitalia offers the most diagnostic features, particularly the structure of the gonostyle. The volsella in New World Pison is usually represented by a pair of lobes associated with the inner margin of the gonostyle. They are often quite large as in the cressoni group (fig. 145), but in the pilosum group the volsellar lobes are much reduced (fig. 308) and sometimes scarcely defined. See species group discussions for further details. CHARACTER ANAL VSIS Discussions of characters and their polarity in the subfamily Larrinae were given by Bohart and Menke (1976) and Lomholdt (1985). Lomholdt presented a cladistic analysis and a reclassification of the Larrinae. The following review of characters is largely restricted to those features that are of significance in Pison and not polarized by Bohart and Menke (1976:224). In my attempts at polarizing characters I have studied examples of most genera in all subfamilies of the Sphecidae, but I have examined as many species of each genus in the Trypoxylini as possible. I have also reviewed Brothers' (1975) important paper on aculeate phylogeny, as well as Carpenter (1981) on the Vespidae. Determining polarity has sometimes been problematical and developing a hypothesis of mandibular notch polarity has been especially unsatisfying. What is really needed is a thorough morphological study of the mandible throughout the Sphecidae similar to the one done on bees by Michener and Fraser (1978) in which all grooves, ridges, setal rows, etc. are analyzed and homologies developed. The plesiomorphic state for each character is listed first. Mandibular notch: 1. absent. 2. present. There can be scarcely any doubt that the notched mandible is an apomorphy within the Sphecidae as a whole, but how to interpret it within the subfamily Larrinae, where it is a common feature, is a problem. Lomholdt (1985:19, 21)
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regarded a notchless mandible as the plesiomorphic condition in the Larrinae ~nsu lato, within which he incorporated the Crabroninae (Oxybelini and Crabronini). He stated that notchless mandibles occur in "a very few larrine genera", but unfortunately he did not list them. Bohart and Menke (1976:225) declined to polarize the presence of a mandibular notch, but suggested that the notch could be the plesiomorphic state. The notch is universal, or nearly so, in the majority of the 39 genera in the Larrinae sensu stricto (Crabroninae excluded), and is absent primarily in genera that are fairly specialized (Nitela, Auchenophorus, Aulacophilus, Aulacophilinus, Pisoxylon, Trypoxylon, Sanaviron, Bohartella, Oalara and Paraliris). Lomholdt (1985: 19, 45) considered the simple mandible in Nitela and Auchenophorus to be a reversal from the apomorphic notched state, and presumably that was his interpretation of the presence of simple mandibles in the 30 or so genera of the Crabronini. Regardless of the polarization adopted, homoplasy is involved, and support for either hypothesis can only be obtained by a thorough phylogenetic analysis of all genera in the Larrinae and Crabroninae. In lieu of such an analysis I am adopting the most parsimonious hypothesis of mandibular notch polarity, one that minimizes reversals. Thus I regard a notchless mandible as the plesiomorphic state in Larrinae. I also hypothesize that highly evolved genera like Nitela, Auchenophorus, Trypoxylon, and Bohartella have retained a primitive mandible (instead of a reversal as espoused by Lomholdt). Reversals may have occurred from the apomophic notched condition in genera like Uris, Holotachysphex and Tachytes (Larrini) where a few species have simple mandibles. Notchless mandibles also occur sporadically in a few species of some other genera in the Larrinae §.. §.., and most species of Solierella (Miscophini) have simple mandibles. Bohart and Menke (1976:292) and Lomholdt (1985: 165) believed the notchless mandible in Solierella to be a reversal, but it seems more parsimonious to regard the genus as having retained the plesiomophic notch less mandible in the majority of its species, especially since it is regarded as one of the least specialized genera in the Miscophini by these three authors. A mandibular notch is not unique to the Larrinae §..§.. The condition occurs in eight of the approximately 45 genera of the Crabroninae and also in three small unrelated genera: Dinetus (Astatinae), Laphyragogus (Laphyragoginae), and Xenosphex (Xenosphecinae). Several important apomorphies are shared by the Larrinae and Crabroninae (single midtibial spur, simple volsella, and larvae with preapical anus''). but Bohart and Menke (1976:30, 222) did not combine them into one subfamily. Lomholdt (1985), however, has taken this step and I am in accord. The mandibular notch is an additional derived feature of the Larrinae and Crabroninae Which, although not present in all genera, suggests that it is logical to place the crabronine tribes Oxybelini and Crabronini within a restructured Larrinae. Four of the five genera of Oxybelini have notched mandibles although the state is not universal in all of them. Only four genera of Crabronini have notched mandibles and they were regarded by Bohart and Menke (1976) as among the most primitive in the tribe, a judgement that is probably erroneous due to the presence of various apomorphic states, one, of course, being the mandibular notch. I hypothesize that most genera in the Crabronini have retained the plesiomorphic notchless mandible although a few reversals may have occurred. The function of the notch is unknown although Lomholdt (1985: 19) declared that it and "the row of stiff bristles presumably serve to clean and protect the opposite
*
Lomho1dt (1985:21) regards the absence of larval papi llae as an additional apomorphy, but Evans (1964) treated this state as the p1esiomorphic condition in Sphecidae. Lomho1dt's idea only requires reduction of the papilla along several different phyletic lines, but Evans' po1ari ty requi res reversal s as di scussed by him (Evans, 1964: 232-233): reduct ion of the antenna1 papi lla and then redevelopment of long papi llae "by at least two stocks of Sphecidae". In bees Michener (1953) regarded the absence of papillae as an apomorphy. More study of this character seems appropriate. Lomho1dt also introduced a new apomorphy for the Larrinae, the presence of a "genal tentorial arm", but he added that "the significance of thi s character can be evaluated better after studyi ng more sphecid genera". He also used a carinate hindcoxa as a synapomorphy for the Larrinae but see my discussion on page 17.
Menke: Pison in the New World
15
mandible". How a mandibular notch can clean any part of the opposite mandible escapes me, and it is more likely that the "bristles" are used in digging, Le., part of the psammophore. It is conceivable that the notch might be used in cleaning an antenna or part of a leg, but no one has described such activity to my knowledge. The notch is common in fossorial genera and it may have some function in nest building, although the fact that some fossorial genera in the Crabronini have notchless mandibles confounds this idea. The absence of a mandibular notch sometimes coincides with a xylicolous or mud nest building lifestyle as in Trypoxylon. Some Pison are ground nesters (chilense, several Australian species - see Evans, 1981) and they have notchless mandibles. Members of the convexifrons and pilosum groups of Pison are nonfossorial so far as known, and all of them have notched mandibles. The small genus Holotachysphex is also perplexing; five of its six species have notchless mandibles, but the one with notched mandibles and the only species whose biology is known, turneri (Arnold), is a twig nester (Gess, 1978). In the final analysis, there is no strong evidence for associating the mandibular notch with fossorial or nonfossorial habits. Labrum: 1. quadrangular, free margin entire. 2. bilobed. -----=rhe labrum is a fairly simple structure in Pisonopsis, Aulacophilus and many species of Pison, but the apex is variably emarginate in some Pison and Trypoxylon. The extreme elaboration of the emargination results in a labrum that has two long, finger-like lobes (Pisoxylon, some Pison, some Trypoxylon), an obvious apomorphy. Anterior polished rim of pronotum: 1. narrow. 2. broadened. Generally the anterior rim of the pronotum in the Sphecidae is narrowly polished, presumably as a bearing surface for the back of the head. This is the plesiomorphic condition. This rim is sometimes elaborated into a mediodorsal triangle as in Prionyx (Sphecinae), or a broad dorsal boss as in some Pison. Sometimes the rim is broadened over much of its length as in Ampulex (Ampulicinae), Polemistus and Arpactophilus (Pemphredoninae), Psammaletes and Ammatomus (Gorytini), Paranysson and especially Mesopalarus (Miscophini), or it may be broadened laterally as in Vechtia (Crabronini). These various elaborations are obvious apomorphies. Anterodorsal lamella of pronotum: 1. absent. 2. present. In some Pison the narrow polished rim is elaborated into a broad, free lamella, an obvious apomorphy. Within the Trypoxylini this lamella is not unique to Pison. It occurs in Aulacophilus and some Trypoxylon (rugifrons for example). In a cursory survey of the other sphecid groups I have found a similar structure in the Sphecinae (Trigonopsis, Podium), the Pemphredonini (Pemphredon, Stigmus), the Scapheutini (Scapheutes, Bohartella), the Crabronini (Entomocrabro, Quexua, Chi mila) and the Gorytini (Sagenista, Pseudoplisus oraclensis). A similar but possibly not homologous lamelliform structure occurs in some Pison and elsewhere in the Sphecidae (Neodasyproctus, Dasyproctus and EctemniLiSOf the Crabronini for example). The anterodorsal pronotal pit or transverse depression is often delimited cephalad by a carina which, in some taxa, is elaborated into a lamella that resembles the polished anterodorsal lamella, but it is dull, setose and has at its base the normal, narrow polished, pronotal rim. The presence of the latter suggests that this type of lamella is not homologous with the polished lamelliform extension of the pronotal rim. In any event, both forms of lamellae are obvious apomorphies. Anterodorsal pit of pronotum: 1. absent 2. present. The anterior part of the pronotum of many sphecids has a pit, a pair of pits, a series of pits, or a transverse depression that may be sulciform. These are believed to be homologous but some of the linear depressions have a more anterior location and may represent independent developments. Pro notal depressions are absent in most Ampulicinae, Laphyragoginae, Nyssoninae, Philanthinae, Entomosericinae and Xenosphecinae, and this is the common state in Pompilidae and many Vespidae ~.!. based on my cursory survey of those two groups. On this evidence a plain pronotum is the plesiomorphic condition in Sphecidae (Carpenter and Cumming, 1985, regarded
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similar pronotal pits or "foveae" as apomorphic in Eumeninae). I regard the single, round, median pit seen in some Pison as the simplest apomorphic state. Often the pit is transversely elongate and one or both margins may be cariniform, and this represents further specialization. The channel-like or sulci form depression seen in some sphecids (most Sphecinae, Astatinae, some Pemphredonini, Scapheutini, some Crabronini, some Trypoxylini) may be a further extension of this trend, or else a completely independent specialization. In genera having species with a pronotal lamella, a polished linear depression is sometimes closely associated and it is likewise possibly a completely independent development instead of a highly derived pit. Its location is more forward than the normal location of the pit. I have made a cursory examination of the pronotum throughout the Sphecidae and present my findings below. In the Trypoxylini I have examined many species of each genus, but only one or two species of each genus in other sphecid groups. My survey of the genera in the Pemphredoninae and Nyssoninae has been even less thorough. Thus. the following discussion is preliminary, but it does give some indication of the variation of a new character complex. In the Trypoxylini only Pisonopsis, Aulacophilus, and Pison have depressions (I have not seen material of Aulacophilinus Lomholdt, 1980). I examined a cross section of species in Trypoxylon (including Trypargilum) and the pronotum always lacks a pit or transverse depression. The same is true of Pisoxylon. Although most species of Pison have a pronotal pit or transverse depression, the plesiomorphic pitless condition is occasionally seen (obliteratum, xanthopus, multistrigatum and kohlii in the Old World; species of the euryops, krombeini and pilosum groups in the New World). In the Miscophini the pronotum is pit less in Nitela, Auchenophorus, Mesopalarus, some Solierella, some Plenoculus, some Miscophus, some Saliostethus and in Nanomiscophus. Lyroda is unique in having a linear series of large, deep pits that closely parallel the polished rim of the pronotum - an autapomorphy for the genus. The forward location of these pits may mean that they are not a homologue of the pronotal pit. I suspect they are not. Other genera in the Miscophini have a transversely elongate pronotal pit (Plenoculus, Sphodrotes, Larrisson, Paranysson, Aha and some Solierella and Miscophus). Some Saliostethus have two pits. I have not examined Sericophorus or some of the endemic African genera. In the Oxybelini and Crabronini the pronotum may be pit less, or more commonly have a transverse depression that is sometimes vaguely defined. In some genera there are two discrete pits (Enoplolindenius, Foxita, Arnoldita, Chimiloides, Lestica, for example). Anacrabro and Encopognathus have a deep, transverse groove. In the Bothynostethini the pronotum is plain in all species of Bothynostethus examined, and Willinikiella argentina (Schrottky) only has a vague pronotal depression. I have not examined Sanaviron (Vardy, 1987). Most genera in the Larrini seem to have a pair of elongate pits or a single elongate pit that may be weakly impressed. Some species of Tachysphex have a small circular pit. In Palarus (Palarini) the pronotum has a pair of transversely elongate grooves that are narrowly separated at the midline by a bridge. In some species there is a transverse groove behind these two depressions that is similar to the condition found in some nyssonine wasps (see below). In the Pemphredoninae the pronotum displays a variety of states. The Psenini that were examined have a transverse impression at the base of the collar, but it is often vague, and it is not clear if this is homologous with the pronotal pit. In the Pemphredonini the pronotum may be pitless (Diodontus, Polemistus), have a transverse channel that may be deep (Pemphredon, Stigmus) or shallow (Carinostigmus), or a pair of shallow pits (Spilomena, Ammoplanops, Pulverro). The front margin of the pronotum is deeply emarginate at the middle in the last two genera, a feature that may have taxonomic importance. Although most nyssonine taxa seem to have a plain pronotum, Heliocausus (including Tiguipa and Acanthocausus - Heliocausini) has a transverse depression at the base of the collar. Similar depressions occur in the gory tin genera Sphecius, Handlirschia and Kohlia; the depression is broadly oval and deep in the last genus. Stizus and Stizoides (Stizini) and some genera in the Bembecini (Hemidula, Stictia, Rubrica, Zyzzyx, and Trichostictia) have a deep pit also, and a thin, sulciform depression often extends laterad from the area. The nyssonine "pit", by virtue of its
Menke: Pison in the New World
17
location at the base of the collar, may not be homologous with the pronotal depressions found in other sphecid subfamilies. Although a pitless pronotum is regarded as plesiomorphic here, it seems likely to me that reversals have occurred in some sphecid genera. Lateral ridge of propodeum: 1. absent. 2. present. This ridge is a common feature in many genera of the Oxybelini, Crabronini, and Trypoxylini but is infrequent elsewhere in the Larrinae, and the rest of the Sphecidae for that matter. Within the Trypoxylini it is prevalent in Pison and Trypoxylon but not universal in either. Because the Oxybelini and Crabronini are generally regarded as among the most highly evolved larrines, and because the ridge is restricted primarily to the Larrinae, I am considering the presence of this ridge to be an apomorphy. I suspect that reversals may have occurred in Trypoxylon, Pison, and perhaps in other genera, but further study is needed to be sure. Hindcoxal carinae: 1. absent. 2. inner carina present. 3. inner and outer carinae present. Brothers (1975:524, 544) considers a noncarinate hindcoxal dorsum to be plesiomorphic in the Aculeata, and in the Sphecidae. Within the Sphecidae a noncarinate coxa is typical of some species of Ampulex (Ampulicinae), all genera in the Sphecinae, a few genera in the Pemphredoninae, all genera in the Astatinae, the Laphyragoginae, all Nyssoninae except Nyssonini, some Alyssonini and some Gorytini, and all Philanthinae except Pseudoscolia and some Cerceris. According to Lomholdt (1985:20) the hindcoxa is carinate "in most larrine genera" and "a very similar structure is present in Dinetus" (Astatinae). But in fact the hindcoxa is noncarinate in all genera in the Larrini (rare exceptions), some species of Palarus (Palarini), some Trypoxylini (Pisoxylon, some species of the subgenus Trypargilum, and rarely in Pison), the genus Belomicroides (Oxybelini), approximately half of the genera in Crabronini, and some genera of Miscophini (some Lyroda, apparently all Plenoculus, a few Miscophus, apparently all Sphodrotes, many Paranysson, and all Aha). Finally, three species of Dinetus available to me, pictus (Fabricius), cereolus Morice, and psammophilus Kazenas, lack hindcoxal carinae. When a carina is present it is typically found along the inner edge of the dorsum. Sometimes it is lamelliform and may be extruded into a tooth near the coxal base. In some sphecids, there are two carinae, an inner one and an outer one (Entomosericus, Bohartella, Enchemicrum, Oxybelus, Encopognathus, Entomognathus, Lindenius, some Palarus, some Larrisson, some Auchenophorus, some Pison, most Pisonopsis). The outer one is often restricted to the apical one-half of the coxa. In rare cases, the outer carina is the only one present (Belomicrus, ~hirflila, some Lestica for example). The hindcoxal carinae vary in form and length in different sphecid groups, and I suspect that these carinae have arisen independently a number of times; the possibility of a few reversals also needs to be examined. Carpenter (1981) regarded the hindcoxal carina as homoplasious in the Vespidae. Lomholdt (1985:21) cites the carinate hindcoxa as one of five synapomorphies for the subfamily Larrinae, adding that "no such carina occurs elsewhere in the sphecid wasps". But this in incorrect. My cursory survey of the Sphecidae indicates coxal carinae occur in several other subfamilies. In the Ampulicinae apparently all members of the Dolichurini have a coxal carina and so do some species of Ampulex (Ampulicini). I have not surveyed the Pemphredoninae thoroughly but a carina is present in Ammopsen, Stigmus, Ammoplanops, Pulverro, and some Pemphredon. Apparently all genera in the Nyssonini and the gory tin genera Clitemnestra, Ochleroptera, Olgia and Argogorytes (some) have coxal carinae. A short carina is found in some species in the Alyssonini, and some species of the philanthine genera Cerceris and Pseudoscolia. In summary, a hindcoxal carina is a common feature in the Larrinae, but it is not universally present nor is it unique to the group by any means. Thus, it cannot be a synapomorphy for the Larrinae.
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Tarsal claws: 1. symmetrical, gradually attentuate to apex. 2. asymmetrical or suddenly attentuate near apex. In Pison the claws of each leg are nearly always identical and typically they narrow gradually toward the apex. In the pilosum group, however, the anterior claw of the foreleg and sometimes the mid leg is deformed in the male, an apomorphy. Another apparent apomorphy is the peculiar thick claw found in some species where attenuation to the apex occurs suddenly (delicatum. agile, stangei and krombeini groups). Male sternum VIII: 1. apex not emarginate. 2. apex with semicircular or V-shaped emargination. 3. emargination bounded by a pair of blunt projections. The emargination of sternum VIII results in the presence of a pseudosting that may be elaborated upon by the development of small projecting lobes. I consider these to be derived traits. Gonostyle of male genitalia: 1. narrowly elongate distally. 2. with short distal lobe. An elongate, setose gonostyle is the common condition in Larrinae, and most Pison, but in the pilosum group the gonostyle is often represented by a large plate that has at most a short apical setose lobe. Gonostylar appendages: 1. none present. 2. one accessory lobe present ventrally. 3. two accessory lobes present ventrally. A fairly simple gonostyle was considered by Bohart and Menke (1976) as the plesiomorphic state. A few New World species of Pison have elaborate gonostyli including one or two lobes ventrally. Volsellar lobes: 1. large. 2. small Reduction of the volsella in the Sphecidae is generally accepted as the apomorphic condition (Bohart and Menke, 1976; Lomholdt, 1985). Within the Larrinae the volsella is sometimes absent (Paranysson, Plenoculus, Solierella, Nitela - all in Miscophini) or reduced to a small setose area or lobe on the inner basoventral area of the gonostyle (Aha, Miscophus, Saliostethus, Saliostethoides, Miscophoides, Namiscophus - all Miscophini). The majority of larrine genera have a well defined volsella, however, and this is true for most species of New World Pison. But the volsellar lobes are much reduced in the pilosum group. The elongate volsellar lobes of the cressoni and eremnon groups may be elaborations, so that reversals can not be ruled out. BIOLOGY Data are available for only 7 of the 44 species treated here and much of it is fragmentary so that no generalizations can be made. Nothing is known for 7 of the 12 species groups recognized. Pison chilense excavates burrows in the ground and cells are separated by mud partitions. The other species, agile, cressoni, aranevorax, longicorne, pilosum and sphaerophallus, make nests in sheltered situations such as bird nests, wasp nests, tents of lasiocampid moths, and the underside of leaves. These non-groundnesters make mud cells that are sometimes arranged in rows, sometimes randomly grouped. Spiders are provisioned. Chalcidoid parasitoids have been reared from the nests of some species. See individual accounts of the above species for further details.
Menke: Pison in the New World
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KEY TO SPECIES OF NEW WORLO PISON" (unknown: males of abathes, duckei, erebus, eyvae, martini, nosferatu, neotropicum, pentafasciatum, and phthinylla, and females of lillo and oaxaca) I.
Mandible with posterobasal notch (figs. 5-6) ................................................... 2 Mandible entire, without posterobasal notch (fig. 3) ....................................... 19
2.
Females (six gastral segments) ....................................................................... 3 Males (seven gastral segments) ..................................................................... 10
3.
Anterior margin of pronotum with broad, non-Iamelliform, polished area that is restricted to middorsum (fig. 288) ............................................................... 4 Anterior margin of pronotum with broad, polished, free lamella that extends laterad to level of pronotal lobe (figs. 294-295) ........................................... 7
4.
Flagellomeres VII-IX nearly three times as long as wide; clypeal lobe as in fig. 285; propodeal hindface unridged (or largely so) .... longicorne Menke, p. 78 Flagellomeres VII-IX about as long as wide; clypeallobe as in fig. 270; propodeal hindface with transverse ridges .................................................................. 5
5.
Propodeal dorsum finely, transversely carinate or striatopunctate (fig. 268) .................................................................... convexifrons Taschenberg, p. 75 Propodeal dorsum smooth except for median carina (fig. 269) ........................... 6
6.
Mesopleural punctation sparser toward mesopleural sulcus (fig. 280); Amazon basin and northward ...................................................... cooperi Menke, p. 77 Mesopleural punctation uniformly dense to mesopleural sulcus (fig. 279); northwestern Argentina ............................................. wasbaueri Menke, p. 76
7.
Gaster with obvious constriction between terga I-II in lateral profile (fig. 301), tergum I strongly humped ..................... (pilosum complex) .......................... 8 Gaster without constriction, tergum I not humped ........................................... 9
8.
Antenna completely black .................. pilosum Smith & vincenti Menke, p. 81, 84 Flagellomeres I-II pale beneath; se Brasil, Paraguay .... aureofaciale Strand, p. 84
9.
Propodeal dorsum with median carina and lateral line of foveae; clypeal lobe arcuate (fig. 324); frons with ocellar-sized polished depression (fig. 323) .................................................................................... gnythos Menke, p. 86 Propodeal dorsum without median carina or lateral line of foveae (fig. 337); clypeallobe triangular (fig. 333); frons without ocellar-sized polished depression above sockets ..................................... sphaerophallus Menke, p. 88
10. Foretrochanter with sharp spine on underside (figs. 292-293) .......................... II Foretrochanter without spine on underside .................................................... 14 II. Foretrochanter spine located near middle of segment (fig. 293); propodeal dorsum without median longitudinal carina; propodeum without crenulate groove laterally ............................................................. sphaerophallus Menke, p. 88 Foretrochanter spine located at apical third of segment (fig. 292); propodeal dorsum with median, longitudinal carina, at least basally; propodeum with crenulate groove laterally ..................... (pilosum complex) ....................... 12 .. Head measurements should be made at a magnification of at least SOX.
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12. Sternum VII flat, without circular or oval depression at middle (fig. 303); anterior claw of midleg normal, not distorted ............................... pilosum Smith, p. 81 Sternum VII with circular or oval depression at middle (may be concealed under S VI) (figs. 304-306); anterior claw of midleg distorted ............................... 13 13. Sternum VIII narrow and depression on VII circular (fig. 304); s. Brasil, Paraguay ............................................................................. aureofaciale Strand, p. 84 Sternum VIII broad and depression on VII elongate (fig. 305); n. South America ................................................................................... vincenti Menke, p. 84 14. Sternum VIII rounded apically (figs. 306, 328) ................................................ 15 Sternum VIII emarginate apically (fig. 278) .................................................... 16 15. Tergum VII compressed laterally, forming a median ridge; frons without ocellus-sized polished spot between sockets and midocellus; UID more than one-half LID ................................................................. oaxaca Menke, p. 85 Tergum VII without median ridge, apical margin reflexed; frons with polished ocellus-sized median spot between sockets and midocellus; UID less than one half LID ....................................................................... gnythos Menke, p. 86 16. Flagellomeres VIII-X almost twice as long as broad; propodeal hindface smooth or nearly so .................................................................. longicorne Menke, p. 78 Flagellomeres VIII-X about as long as broad, shorter and thicker than I-III; propodeal hindface with strong transverse ridging (figs. 268-269) ................ 17 17. Propodeal dorsum cross-carinate (fig. 268) ......... convexifrons Taschenberg, p. 75 Propodeal dorsum largely smooth (fig. 269) ................................................... 18 18. Gonostyle of genitalia with long, curving spine ventrally (fig. 282); Bolivia, Brasil north to Costa Rica, Dominica ....................................... cooperi Menke, p. 77 Gonostyle with short ventral spine (figs. 276-277); northern Argentina ................................................................................. wasbaueri Menke, p. 76 19.
Eye covered densely with short setae (fig. 9) ................................................. 20 Eye bare or with only few widely scattered setae .......................................... 26
20. Propodeal side without dorsal carina or ridge; forewing with two or three submarginal cells ..................................................................................... 21 Propodeal side delimited dorsad by carina or crenulate ridge that extends from petiole socket area to spiracle (carina may be obscured by setae); forewing with two submarginal cells ......................... (stangei group) ....................... 23 21. Forewing with two submarginal cells, the second not petiolate; tergum I simple apically, not reflexed nor with subapical transverse depression; northeastern North America ................... (agile group) ......................... agile (Smith), p. 38 Forewing with three submarginal cells, the second petiolate; tergum I with reflexed apical rim (with subapical transverse depression); Neotropical .......................................... (krombeini group) .......................................... 22 22. Propodeal dorsum and hindface densely covered with appressed silver or gold setae that obscure shiny surface; sternum I with appressed silver setae; disk of scutum shiny, parapsidal and admedian lines sulci form ................................................................................. krombeini Menke, p. 45 Propodeal dorsum with dense appressed setae at base, but apex and hindface sparsely setose, not obscuring dull surface; sternum I with erect pale setae (not silver); scutum dull, parapsidal and admedian lines not sulci form ............................................................................. neotropicum Menke, p. 46
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21
23. Ocellocular distance 1.38 to l.65X hindocellus diameter; propodeal dorsum smooth. punctate; eastern and southeastern Brasil ...... plaumanni Menke, p. 40 Ocellocular distance l.08 to 1.31 X hindocellus diameter; propodeal dorsum variable .................................................................................................. 24 24. Upper interocular distance. 76 to .80X the distance between eye notches, the latter shallow (fig. 78) ................................................... abathes Menke. p. 43 Upper interocular distance .59 to .66X the distance between eyenotches; eye notches deep (fig. 9) .............................................................. 25 25. Pronotum rounded laterally, humerus at most with faint ridge (fig. 94); propodeal dorsum mostly ridged or striatopunctate; occipital carina complete ventrad; Bolivia. Argentina ......................................................... stangei Menke, p. 41 Pronotum angulate laterally, humerus with obvious ridge (fig. 98); propodeal dorsum mostly punctate (ridging restricted to base and short cross-carinulae associated with median carina); occipital carina interrupted ventrad; Panama to central Brasil ............................................................ duckei Menke, p. 42 26. Propodeal side delimited dorsad by carina or crenulate ridge that extends from petiole socket area toward spiracle (figs. 27, 46) ........................................ 27 Propodeal side without dorsal carina or ridge ................................................ 39 27. Eye length 10-20% less than distance between eye notches (fig. 10); propodeal dorsum without median longitudinal carina, surface smooth, punctate; labrum ending in two fingerlike lobes .................... (euryops group) ....................... 28 Eye length equal to or greater than distance between eye notches (fig. 15) (if 10% less then propodeal dorsum with median carina); propodeal dorsum with or without median longitudinal carina; labrum arcuate, or truncate or shallowly emarginate apically ................................................................................. 34 28. Females, gaster with 6 visible terga ............................................................. 29 Males, gaster with 7 visible terga ................................................................. 31 29. Forewing with two submarginal cells, outer veinlet of submarginal cell I angled (fig. II); clypeallobe quadrangular, lower lip roundly triangular (figs. 124-125) ............................................................................ styphopteron Menke, p. 50 Forewing with three submarginal cells; clypeus various .................................. 30 30. Propodeal hindface with coarse, transverse ridges from top to bottom; upper interocular distance essentially equal to lower interocular distance .................................................................................... euryops Menke, p. 48 Propodeal hindface smooth above, ridges present only next to petiole socket; upper interocular distance at most .8X as long as lower interocular distance ............................................................................................ eu Menke, p. 47 31. Forewing with two submarginal cells, outer vein let of submarginal cell I angled (fig. 11); flagellum without linear tyli or asetose, shiny areas ............................................................................ styphopteron Menke, p. 50 Forewing with three submarginal cells (if only two then outer veinlet of I is straight); flagellomeres with asetose polished areas or linear tyli present on III-V ....................................................................................................... 32 32. Propodeal hindface smooth above, with transverse ridges only near petiole socket ............................................................................................ eu Menke, p. 47 Propodeal hindface with coarse transverse ridges from top to bottom ............. 33 33. Flagellomeres V-VI asymmetrically swollen in profile (fig. 126) .................................................................................... euryops Menke, p. 48 Flagellomeres V-VI not swollen .............................................. lillo Menke, p. 49
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34. Occipital carina a complete circle, separated from hypostomal carina by at least an ocellus diameter; pronotum with broad, transverse lamella anterodorsally (fig. 43) ...................... (fritzi group) ........................................................ 35 Occipital carina incomplete ventrally. or contiguous with hypostomal carina; pronotum with or without anterodorsal lamella .......................................... 36 35. Carinae of propodeal dorsum mostly transverse, giving way to punctures laterally (fig. 47); anterior margin of pronotum with broad lamella laterally at level of pronotal lobe (fig. 45) ....................................................... fritzi Menke, p. 31 Carinae of propodeal dorsum oriented posterolaterally, extending nearly to lateral ridge (fig. 49); anterior margin of pronotum without lamella at level of pronotal lobe ............................................................. nosferatu Menke, p. 32 36. Tarsomeres without conspicuous stout setae distally (fig. 71); episternal sulcus ending ventrad without curving forward to anterior margin of mesopleuron; propodeal dorsum with median longitUdinal carina but smooth overall, appearing impunctate ................................................ delicatum Menke, p. 35 Tarsomeres armed distally with conspicuous stout setae; episternal sulcus curving forward ventrally, reaching anterior margin of mesopleuron; propodeal dorsum punctate and/or covered by carinae ................ (conforme group) ................ 37 37. Disk of propodeal dorsum smooth, punctate, shiny. with median longitudinal depression that contains carina (fig. 25) ........................ con forme Smith, p. 25 Disk of propodeal dorsum obliquely carinate or striatopunctate, with or without median longitudinal depression and carina (figs. 28-29) ............................... 38 38. Propodeal side almost entirely smooth, punctate; Mexico? ................................................................................. doggonum Menke, p. 26 Propodeal side striatopunctate on dorsal half (fig. 26); Ecuador to Bolivia ............................................................................. some larsoni Menke, p. 29 39'. Tergum I sharply depressed subapically, forming a broad band; propodeal side with a few large punctures scattered among dense, fine punctation (fig. 57); body 12.5 mm long or more .......................................... eremnon Menke, p. 33 Tergum I without sharp subapical depression; punctures of propodeal side of uniform size; body 12 mm or less .............................................................. 40 40. Propodeal dorsum with many carinae, at least basally (figs. 29, 31, 263), or striatopunctate ........................................................................................ 41 Propodeal dorsum at most with median, longitudinal carina, surface smooth punctate .............................. (cressoni group*) ......................................... 44 41. Pronotal collar thin, knife-edged, closely appressed to and lower than scutum; appressed silver setae confined to lower frons ........ (con forme group) ......... 42 Pronotal collar as thick as metanotum, rounded, and nearly as high as scutum; head and thorax with extensive appressed silver setae .... (chilense group) .... 43 42. Ridges of propodeal dorsum obliquely oriented (fig. 29) ............................................................................. some larsoni Menke, p. 29 Ridges of propodeal dorsum longitudinally oriented when present (fig. 31) ....................................................................................... eyvae Menke, p. 27
* Most species in this group are separated reliably only by rather subtle differences in the female clypeus and male genitalia. I have attempted to use other, more obvious differences in the key (color for example), realizing that they may not work for all material. Comparison with descriptions and figures should be made after arriving at a name in the key.
Menke: Pison in the New World
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43. Wing veins amber; erect setae of head, thorax and legs black; terga I-III with bright silver fasciae; propodeal dorsum obliquely carinate/striatopunctate .................................................................................. chilense Spinola, p. 71 Wing veins black; erect setae white; tergal fasciae dull; propodeal dorsum longitudinally carinate, interspaces impunctate ................. ~ Menke, p. 73 44. Forewing with two submarginal cells (fig. 132) .............................................. 45 Forewing with three submarginal cells (fig. 131) ............................................ 47 45. Legs yellowish brown except coxae; first segment of gaster almost entirely yellowish brown; female clypeus as in fig. 249; female tergum II strongly humped (fig. 251) ....................................................... phthinylla Menke, p. 70 Legs largely or completely black; gastral segment I black except pale apical band sometimes present on tergum; female clypeus as in figs. 160, 239, 241; female tergum II not humped ............................................................................... 46 46. Ocellocular distance .25 to .40X hindocellus diameter; female clypeus as in figs. 239, 241 ........................................................... aranevorax Menke, p. 68 Ocellocular distance. 71 X hindocellus diameter; female clypeus as in fig. 160 ...................................................................................... ere bus Menke, p. 57 47. Legs entirely yellowish brown (coxae and hindfemur above sometimes black) ... 48 Legs black, or, at most, femora and tibiae a mixture of black and pale brown .. 50 48. Forewing media diverging from M+Cu before crossvein cu-a; female clypeus as in fig. 230; male clypeus as in fig. 231 ..................... abothrum Menke, p. 67 Forewing media diverging from M+Cu after cu-a ........................................... 49 49. Gastral segment I all black, no yellow apical band on tergum; tergum II strongly convex in lateral profile (fig. 162); tergum I longer than wide .................................................................................... martini Menke. p. 58 Gastral segment I completely or extensively yellowish brown, tergum with yellow apical band; tergum II not unusually swollen; tergum I as long as wide .............................................................................. maculipenne Smith, p. 59 50. Scutellum more sparsely punctate posteriorly where very fine punctures are mixed with large punctures (fig. 142); female clypeallobe reflexed (fig. 136) .................................................................................. cressoni Rohwer, p. 53 Scutellum uniformly punctate, punctures of one size ...................................... 51 51. Upper interocular distance .86-.91 X lower interocular distance in female, .92-1.0X in male ...................................................................................... 52 Upper interocular distance .57-.79X lower interocular distance in female • . 69-.86X in male ...................................................................................... 53 52. Female clypeus as in fig. 177; male flagellum with linear tyli at least on articles IV-VI; forewing media diverging from M+Cu before crossvein cu-a or interstitial with it ...................................................... brasilium Menke, p. 61 Female clypeus as in fig. 221; male flagellum without tyli; forewing media diverging after cu-a or interstitial with it .................... dementia Menke, p. 66 53. Propodeum largely covered by dense gold setae that obscure surface and ocellocular distance in female usually .82-1.16X hindocellus diameter, and in male .93-1.18X HOD; female clypeus as in figs. 149-151 ................................................................................... chrysops Menke, p. 55 Propodeal surface largely or entirely visible, but if dense setae present on dorsum then ocellocular distance in female no more than .69X hindocellus diameter and in male no more than .85X HOD; female clypeus different ...... 54
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54. Ocellocular distance 1.23X HOD; terga I-V with broad, yellow apical bands ......................................................................... pentafasciatum Menke, p. 58 Ocellocular distance .33 to .85X HOD; yellow bands confined to terga I-III ...... 55 55. Female clypeus as in figs. 204, 206, 209; male flagellum with linear tyli on articles III-V or VI (fig. 213) .......................................... cameronii Kohl, p. 64 Female clypeus as in figs. 166, 168-169, 186, 189, 192, 194; male flagellum without tyli ............................................................................................. 56 56. Female clypeus with short median lobe whose margin is straight or slightly sinuate (figs. 166, 168-169), clypeal rim not elevated above adjacent punctation; male clypeus as in fig. 170; wing membrane sometimes yellowish .............................................................................. maculipenne Smith, p. 59 Female clypeal margin above labrum with concavity bearing four angles or teeth (figs. 186, 189, 194), clypeal rim elevated above adjacent punctation (figs. 188, 196); male clypeus as in fig. 197; wing membrane never yellowish ........................................................................... arachniraptor Menke, p. 62 Conforme Group (F igs. 15-40) Description: Frons moderately swollen; labrum quadrangular; female mandible with weak mesal tooth on inner margin; male antenna simple, flagellomeres elongate; clypeal disk swollen, punctate; eye length slightly greater than or equal to distance between eye notches (fig. 15); occipital carina ending just before joining apex of hypostomal carina; pronotum without anterodorsal lamella, but with anterodorsal pi t that is sometimes transversely elongate (fig. 16), this depression margined behind by carina, surface behind carina shiny and usually with several ridges; scutal flange of variable width; tegula punctate, setose on basal half, impunctate, smooth beyond; propodeal side sometimes delimited dorsad by crenulate ridge that extends from petiole socket to spiracle (absent in eyvae, sometimes absent in larson i); propleuron sparsely punctate on disk; lower end of episternal sulcus curving forward to anterior margin of mesopleuron (evanscent ventrad in eyvae and females of larsoni); male foretrochanter without spine; outer carina of hindcoxa weak, reduced to distal ridge; hindtarsomere IV with small plantula; male tarsal claws symmetrical; forewing media diverging after crossvein cu-a; forewing marginal cell acuminate apically; three submarginal cells, inner and outer vein lets of III usually broadly separated on marginal cell; hamuli divided into two groups; gaster without yellow bands; tergum I single-edged apically; male sternum VIII rounded or truncate apically, not emarginate; genitalia robust, gonostyle slender and densely fringed with long setae apicoventrally, volsellar lobe large, feebly setose, aedeagus unusually broad (dorsal view, figs. 36, 40), penis valves fused over most of their dorsal length, each with apical horn (figs. 34, 38). Included species: Pison conforme, doggonum, eyvae, and larsoni. Discussion: The main features of this small group are the incomplete occipital carina, the projecting, rounded female clypeal lobe, the coarse cross-ridging of the propodeal hindface, and the non-emarginate male sternum VIII. The dorsolateral propodeal ridge is not universal and can only be termed an apomorphic trend. The globular form of the male genitalia is distinctive but males are unknown in two species. The conforme group does not appear to have a single universal apomorphic character state, and in fact it may be a paraphyletic group. When males of all species are known, perhaps an apomorphy will be found in the genitalia. Pison larsoni and eyvae are sister species, sharing the thin, closely appressed pronotal collar, the absence (except in some larsoni) of a crenulate propodeal ridge.
Menke: Pison in the New World
25
and the evanescing episternal sulcus ventrad in the female. Pison conforme and doggonum have a thicker collar and a well formed crenulate ridge on the propodeum. Pison conforme, by virtue of its transverse pronotal pit and smooth propodeal dorsum, has a somewhat isolated position. Pison doggonum is fairly similar to larsoni in sculpture and general habitus. Pison conforme Smith (Figs. 15-17, 19-20,25,33-36) Pison conforme Smith, 1869:297. Holotype male: "Mexico" (BMNH). Description, female (81 specimens): Black; wings clear or slightly infumate. Frons with appressed silver setae on either side of antennal socket, sometimes dense enough to obscure SCUlpture, clypeus sometimes with similar setae laterally; erect setae on body pale; tergum I with broadly interrupted apical silver fascia. Free margin of labrum with weak indentation. Clypeal lobe as in fig. 19, free margin not thickened or double-edged, surface smooth, polished. Frons dull, shallowly punctate, punctures one to two diameters apart; eye length slightly greater than distance between eye notches (fig. 15); UID 0.47-0.53X LID; ODD 0.33X HOD or less, usually about 0.20X HOD; flagellomere I length slightly less than to slightly more than 3X apical width, II about the same, III-X longer than broad. Anterodorsal pit of pronotum transversely elongate, length equal to UID or greater, carina behind pit sometimes lamelliform (fig. 16); collar thin, laterally with linear impression that makes humeral angle prominent, ridge like (fig. 16); scutum dull or weakly shiny, punctate, punctures usually slightly larger than those of frons and deeper, more sharply defined, separated by less than puncture diameter to two or more diameters (punctures finer than on frons and several diameters apart in specimens from Ecuador), interspaces imbricate (Harris, 1979), hind margin of scutum with short ridges; scutellum shiny or somewhat dull, punctures somewhat smaller than those of scutum; metanotum with micropunctures; disk of propodeal dorsum smooth, shiny, with sparse, fine, setigerous punctures similar in size to those of metanotum, usually with median longitudinal carina (sometimes reduced or absent in specimens from Ecuador) that extends from base to about middle of dorsum (fig. 25), this carina usually in depression (depression absent in specimens from Ecuador), base of dorsum with short perpendicular carinae; propodeal hindface with coarse, well spaced, cross-ridges that are continued onto posterolateral surface of dorsum, interspaces shiny, smooth; propodeal side smooth, shiny, finely punctate, punctures slightly larger, denser than those on propodeal dorsum; mesopleuron shiny, punctate, punctures same size as those on scutum and less than diameter to several diameters apart; metapleuron shiny, with pinprick punctures. First recurrent vein usually ending near middle of second submarginal cell, rarely interstitial between I-II, second recurrent vein ending on submarginal cell 1II or interstitial between II-III. Gaster shiny, terga I-II finely punctate, punctures several diameters apart, sternum II swollen in profile, often with vague gibbosity on midline at apical third. Length 6.5-8.5 mm. Male (33 specimens): Same as female except: clypeus usually covered by dense, appressed silver setae; clypeal lobe as in fig. 20; UID 0.55-0.62X LID; ODD usually 0.25X HOD; scutum usually shiny; first recurrent vein usually ending near base of second submarginal cell, and second recurrent interstitial between II-III, but variety of other arrangements observed; apical margin of tergum VII arcuate; sternum II sometimes swollen, sometimes with vague gibbosity; sterna II-VI each with one or more long, erect setae posterolaterally; sternum VIII apex roundly truncate, slightly indented' (fig. 17); genitalia as in figs. 33-36, aedeagus slightly curved in lateral profile, apical horns not sharply pointed (fig. 34); length 5.5-7.5 mm.
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Discussion: Pison conforme is commonly collected in Central America and northwestern South America. Within the conforme group the smooth. punctate propodeal dorsum with a median longitudinal carina usually set in a depression is distinctive, although the carina is absent in some Ecuadorian material. The transversely elongate pronotal pit. not easily seen unless the head is tipped forward. distinguishes con forme from larsoni and doggonum. Tergum VII is rounded apically in males of conforme. but in larsoni it is truncate. The aedeagus also differs between these two species (compare figs. 33-36 & 37-40). I have studied Smith's type. Range: West central Mexico to Peru. Material examined: MEXICO. Jalisco: 9 mi. s Guadalajara (MCZ); Morelos: Cuernavaca (CU. USNM); Vera Cruz: Jalapa (UCD. USNM); Rio Blanco (UCD); Rinconada (UCD); Fortin de las Flores (FSDA); Orizaba (UCD); Cordoba (GENEVA); Chiapas: San Cristobal de las Casas (CU. CNC); Bochil (UCD); Nachic (CU); L. Montebello N. P. (CNC). GUA TEMALA: Sta. Emilia. Pochuta (USNM); San Jeronimo B. V. P. (AMNH); Helvetia. San Sebastian (MCZ); Las Sabanetas. Barbarena (MCZ); Moca. Guatalon (MCZ). EL SALVADOR: 3 mi. w Quezaltepeque (UCD); Mt. San Salvador (UCD). COST A RICA: Turrialba (BERLIN); Santa Ana (MCZ); San Jose, San Antonio de Escazu (COOPER); San Pedro de Montes de Oca (USNM); San Jose (USNM. BMNH); Alajuela. Chomogo area (USNM); Monteverde (USU); Escazu (USU). COLOMBIA. Valle: Peiias Blancas. 10 km w Cali (CSDA. FSDA); Candelaria. Finca San Luis (CSDA); Pance CVC. 15 km w Cali (CSDA); Cali (BMNH); Boyaca: Muzo (MCZ); Cauca: Chisquio El Tambo (OSU); San Andres de Pisimbala. 60 km e Popayan (BMNH); Magdalena: N. Sierra Nevada de S.• Marta, Rio Buritaca (BMNH); Vista Nieve. San Lorenzo Mt. (USNM); Putumayo: Mocoa (BMNH); Cundinamarca: 3 km n Alban (AMNH). ECUADOR. Pinchincha: Tinalandia near Sto. Domingo (PMA). 47 km s Santo Domingo. Rio Palenque Sta. (CNC); Prov.?: Naranjapata (BPBM); Huigra (CU). PERU. Madre de Dios: Avispas (MCZ). Pison doggonum Menke. n. sp. (Figs. 24. 27-28) Description, holotype female: Black; wings clear. Eye notches and lower frons with appressed silver setae that obscure sculpture. clypeus more sparsely covered by appressed silver setae; rest of head and thoracic vestiture pale or silvery; tergum I with silver fascia laterally, adjacent area of tergum II with some appressed silver setae. Free margin of labrum straight. Clypeal lobe as in fig. 24. free margin not thickened or double-edged, surface smooth. not highly polished. Frons dull. shallowly punctate. punctures one to two diameters apart; eye length slightly greater (l.03X) than distance between eye notches; UID 0.47X LID; OOD 0.06X HOD; flagellomere I length 3X apical width. II almost as long. III-X longer than broad. Anterodorsal pit of pronotum oval. greatest width about equal to hindocellus diameter; collar thin, humeral angle not ridgelike; scutum dull, densely punctate. punctures same size as on frons but mostly less than diameter apart. hind margin of scutum with short ridges; scutellum weakly shining. punctation similar to scutum; metanotum more finely punctate than scutellum; disk of propodeal dorsum impunctate. weakly shiny. with median longitudinal carina and many fine, somewhat irregular. oblique carinae that fade posterolaterally where surface becomes punctate (figs. 27-28); propodeal hindface with several widely spaced transverse ridges that become weaker dorsad; propodeal side shiny, smooth (except for a few ridges adjacent to spiracle). with shallow. setigerous punctures that are one to two
Menke: Pison in the New World
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diameters apart; mesopleuron smooth, shiny, punctate, punctures same size as those on scutum but one to three diameters apart; metapleuron microridged above, smooth, shiny, impunctate below. First recurrent vein ending on submarginal cell I on left forewing, interstitial on right forewing, second recurrent vein interstitial on left wing, ending on submarginal cell III on right wing. Gaster shiny, terga with very fine pinprick punctures; sternum II normal, not swollen. Length 7 mm. Male: Unknown. Discussion: Pison doggonum is rather enigmatic since only one specimen is known, but several features separate it from larsoni, its most similar relative: free margin of labrum straight (usually shallowly emarginate in larsoni), hindocellus nearly touching eye (ODD 0.11-0.30X HOD in females of larsoni), pronotal pit as broad as hindocellus (smaller than ocellus in larsoni), SCUlpture of propodeal dorsum fine (coarser in larsoni, compare figs. 27-28, 29-30), and propodeal side nearly all punctate (diagonally striatopunctate on dorsal half in larsoni). Pison doggonum is a more finely punctate wasp than larsoni and the latter species has a thinner pronotal collar. Range: Presumably Mexico. The type specimen is labelled simply, "Mejico, Mus. Drews." ~:
Holotype female: "Mejico" (ZMC). Etymology: The name doggonum is a Latinization of the American slang expression, "dog gone it", in this instance a reflection of my frustration at having only one specimen with poor collecting data. Pison eyvae Menke, n. sp. (Figs. 23, 31-32) Description, holotype female: Black, wings clear. Body with pale erect vestiture (darkest on scutum) that is longest on propodeal hindface, mesopleural venter and underside of coxae and trochanters; lower frons with dense, appressed, tarnished silver setae that obscure sculpture; no tergal fasciae. Labrum quadrangular, margin entire. Clypeus with prominent truncate median lobe with rounded corners, free edge not thickened (fig. 23). Frons dull, granular. shallowly punctate, punctures one to two diameters apart. Eye length 1.04X distance between eye notches; UID 0.48X LID; OOD 0.20X HOD. Flagellomeres I-III each 3X as long as apical width, flagellomere X length 2X apical width. Pronotum with transversely elongate anterodorsal pit that is margined behind by sharp lamelliform carina whose length is less than UID; collar thin, closely appressed to and lower than scutum. Scutal flange broad posterad, strongly reflexed. Scutum and scutellum dull, densely punctate, most punctures less than diameter apart. Metanotum weakly shiny, densely micropunctate. Propodeum shiny; dorsum densely, finely punctate but with about dozen, short, coarse ridges at base, the two center ridges extending posterad toward apex delimiting linear depression (fig. 32), dorsum laterally with gently arcuate, linear depression that extends from base to near apex; hindface impunctate, with about seven coarse cross-ridges that curve onto propodeal side where they change to fine striatopunctation; propodeal side densely punctate on
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lower half, finely obliquely striatopunctate above but without delimiting ridge. Mesopleuron shiny, densely punctate, punctures one to two diameters apart. Episternal sulcus evanescent as it curves forward ventrad, not clearly reaching pleural margin. Metapleuron shiny, densely micropunctate. First recurrent vein ending at middle of second submarginal cell, second recurrent vein ending on third submarginal cell. Gaster shiny, finely punctate, punctures several diameters apart on terga I-II. Length 9 mm. Variation in females (8 specimens): Wings sometimes weakly infumate (Colombia). Appressed setae of frons sometimes silver (Colombia). Margin of clypeal lobe sometimes arcuate. UID 0.43-0.49X LID (Colombia, Ecuador), or 0.52X LID (Bolivia); OOD 0.17-0.21X HOD (Colombia, Ecuador), or 0.25-0.26X HOD (Bolivia). Carina behind anterodorsal pit of pronotum very high, longer than UID (Bolivia). Propodeal dorsum varies: often only one median, longitudinal carina present (fig. 31), sometimes all basal ridges may be short, i.e, no long median longitudinal carina or carinae, and disk only punctate (Bolivia), or many basal ridges may extend posterad changing to fine, arcuate striatopunctation that covers most of disk with middle ridge strongest and straight (Colombia, fig. 31), lateral arcuate linear depression sometimes absent; propodeal side nearly entirely punctate or irregularly covered with oblique carinulae that are continuations of coarse ridges of hindface (this latter condition occurs in specimens with striatopunctate dorsum). First recurrent vein occasionally interstitial between submarginal cells I-II or ending near base of second submarginal; second recurrent vein sometimes interstitial between submarginals II-III. Length as short as 7.0 mm. Male: Unknown. Discussion: Pison eyvae differs from other members of the conforme group, except larsoni, in having an episternal sulcus that becomes evanscent ventrally as it curves forward toward the pleural margin. Pison eyvae is similar to larsoni in the narrow, closely appressed pronotal collar, female clypeal shape, head measurements, and wing venation, but the sculpture of the propodeal dorsum, though variable, is distinctive in eyvae. The basal ridges are more or less longitudinally oriented, and when long, they curve in a gentle arc toward the apex (figs. 31-32). The uppermost of the lateral, anterad extensions of the coarse ridges on the hindface are similarly curved when viewed from above, thus accentuating the affect. Pison eyvae usually lacks a crenulate ridge at the top of the propodeal side (weakly formed in some specimens), but the ridge is sometimes present in larsoni. Range: Known only from Colombia, Ecuador and Bolivia. ~:
Holotype female: ECUADOR, Morona-Santiago: Macas, 1100 m, II-2-1982, M. Cooper (COOPER). Paratypes (8 females): COLOMBIA, Valle: Lago Calima, 3 mi. behind dam, tropical wet forest, VII-16-75, R. C. Wilkerson (CSDA, FSDA). ECUADOR: same data as type; Pichincha: Nambillo Valley nr. Mindo, 1450 m, VI-26-87, VII-2-87, M. Cooper (COOPER). BOLIVIA, La Paz: Chulumani, 1700 m, IV-3-79, M. Cooper (BMNH), Coroico, 1700 m, V-22-79, M-:-Eooper (BMNH). Etymology: The name eyvae is an arbitrary, euphonious combination of letters.
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Pison larsoni Menke. n. sp. (Figs. 18,21-22,26,29-30,37-40) Description, holotype male: Black; wings slightly infumate. Frons with appressed tarnished silver setae between and lateral to antennal sockets; rest of head and body vestiture pale but generally brownish; no tergal fasciae. Free margin of labrum semicircularly emarginate. Clypeal lobe polished (fig. 22). Frons dull, punctate, punctures separated by one to two diameters, interspaces appearing granulate; eye length equal to distance between eye notches; UID 0.59X LID; ODD 0.36X HOD; flagellomere I length about 2.5X apical width, remaining flagellomeres progressively shorter, VIII-X about as long as broad, flagellomeres II-IV each with a tiny, pale, oval sensory area on ventral side toward base, that on III the largest. Anterodorsal pit of pronotum transversely oval, greatest width shorter than hindocellus diameter; collar thin, closely appressed to and lower than scutum, humeral area not ridge like or otherwise prominent. Scutum dull, punctures same size as those on frons, but somewhat deeper and mostly one diameter apart, interspaces granular; hind margin of scutum with short ridges. Scutellum weakly shiny, more sparsely punctate than scutum. Metanotum densely, finely punctate. Propodeal dorsum shiny, with oblique ridges basally and cross-ridges along center line, these changing to striatopunctation laterally; propodeal hindface with irregular, coarse cross-ridges that become weaker dorsad; propodeal side shiny, obliquely striatopunctate on dorsal half and near hindcoxa, punctate elsewhere (similar to fig. 26). side with irregular crenulate ridge dorsally. Mesopleuron shiny, smooth, punctate, punctures larger than on scutum and about one diameter apart. Metapleuron striatopunctate above, finely punctate below. First recurrent vein of forewing ending on submarginal cell II, second recurrent vein interstitial between submarginal cells II-III. Gaster shiny, terga I-II finely punctate, punctures one to three diameters apart; tergum VI! somewhat drawn out apically, truncate; sterna II-III with one long. erect seta posterolaterally, IV-VI each with two long, erect setae posterolaterally; sternum VIII broad, angularly rounded apically (fig. 18); genitalia as in figs. 37-40, aedeagus strongly arcuate in lateral profile (fig. 37), apical horns acuminate (fig. 38). Length 7.5 mm. Female (7 specimens): -same as male except: free margin of labrum shallowly emarginate or truncate; clypeus as in fig. 21; UID 0.50-0. 54X LID; ODD 0.11-.30X HOD; flagellomere I length 3X apical width, VII-IX longer than wide; scutal punctures mostly half diameter apart; scutellar punctures slightly smaller than those of scutum, less than diameter apart; propodeal dorsum usually with median, longitudinal carina that may be short (figs. 29-30), oblique ridging usually longer, coarser (fig. 29); propodeal side not clearly delimited by crenulate ridge in 5 Ecuador specimens from Nambillo, striatopunctation sometimes covers all of side; episternal sulcus evanscent ventrally as it curves forward; second recurrent vein usually interstitial between submarginal cells II-III, but ending on II in one specimen and on III in another; sternum II with one long, erect seta posterolaterally as in male, but succeeding sterna with more numerous long setae that are more mesal; length 6.5-7.5 mm. Discussion: Pison larsoni has a thin pronotal collar that is lower than and closely appressed to the scutum, a character state shared only with eyvae. The sculpture of the propodeal dorsum, though variable in both species, separates the two: ridges, when present, are primarily longitudinal in eyvae (figs. 31-32), but oblique or transverse and always present in larsoni (at least in females, figs. 29-30). In five of the eight specimens of larsoni the crenulate ridge delimiting the top of the propodeal side is evanescent or absent, a constant characteristic of eyvae. Like eyvae, the female episternal sulcus is evanscent ventrally in larsoni. The striatopunctate upper part of
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the propodeal side of larsoni separates it from doggonum. The ocellocular distance is wider in larsoni than doggonum but too few specimens are available to determine the reliability of this difference. The genitalia of larsoni differs from that of conforme especially in the strongly arched aedeagus and sharp apical horns (compare figs. 37-38 & 33-34). Range: Known only from Ecuador, Peru and Bolivia. ~:
Holotype male: BOLIVIA, La Paz: San Jacinto, 1700 m, XII-S/8-SS, L. Pena (KU). Paratypes (7 females): ECUADOR, Marana-Santiago: Cord. de Cutucu, b km e Macas, 1100 m, V-22-87, M. Cooper (COOPER); Pichincha: Nambillo Valley nr. Minda, 14S0 m, VI-27/28-87, VIII-lS-87. M. Cooper (COOPER). PERU. Cuzco: Valle del Rio Cosnipata. Hacienda Santa Isabel. 1700 m. I-3-S2. F. Woytkowski (ULLO). Etymology: Pison larsoni is dedicated to Gary Larson whose "The Far Side" cartoons have brought much laughter to me and many other entomologists. Fritzi Group (Figs.41-S2) Description: Frons swollen; eye asetose; male antenna with vague sensory areas on some flagella meres; female clypeal lobe trapezoidal or roundly truncate. male clypeal lobe triangular; labrum quadrangular; female mandible with weak mesal tooth on inner margin; occipital carina a complete circle. separated from hypostomal carina by hindocellus diameter or slightly more; pronotum with transversely elongate. polished depression that is margined anteriorly by broad. thin. setose. overhanging lamella; pronotal collar thick but front face flattened broadly at middle; thorax elongate, length in lateral profile about 2X height; scutal flange narrow. upturned; tegula densely punctate on inner half. impunctate beyond; propodeal side delimited dorsad by crenulate ridge that extends from petiole socket to spiracle; propodeal dorsum finely cross-carinate andlor striatopunctate; propleuron punctate; lower end of episternal sulcus curving forward to edge of mesopleuron; outer carina of hindcoxa reduced to short. evanescent. distal ridge; male fore trochanter without spine; plantulae present on tarsomeres II-IV or III-IV; male tarsal claws symmetrical; forewing media diverging after crossvein cu-a; forewing with three submarginal cells; endpoint of recurrent veins highly variable: first to submarginal cell I or II or interstitial, second to submarginal cell II or III or interstitial; gaster without yellow bands; tergum I single-edged apically; male sternum VIII with semicircular emargination apically; genitalia dorsoventrally flattened. penis valve without apical horn. gonostyle densely covered by long. coarse setae ventrally. volsellar lobes feebly setose. Included species: Pison fritzi and nosferatu. Discussion: This small group of rather dull. densely punctate wasps is characterized by the complete occipital.carina that is separated from the hypostomal carina. the dorsal lamella associated with the elongate depression of the pronotum. the thick pronotal collar. the lateral ridge of the propodeum. the apically emarginate male sternum VIII. and the elongate body. The fritzi group is otherwise similar to the conforme group but the male genitalia are robust in the latter and dorsoventrally flattened in the former. Apomorphies are the pronotal lamella. transverse pronotal pit. the propodeal ridge. and emarginate male sternum VIII. The greatly elongate. polished. flat.
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pronotal depression of this group is clearly homologous with the anterodorsal pit of smaller dimensions common in most species of Pison. One species, fritzi, has two lamellae on the pronotum: the one associated with the anterodorsal pit, and one that arises from the front margin of the notum. The last is lamelliform only laterally, narrowing toward the midline of the pronotum. Pison fritzi Menke. n. sp. (Figs. 41, 43. 45-47, 50-52) Description, holotype female: Black; wings clear. Clypeus laterally and lower half of frons except for narrow zone above each antennal socket, densely covered with appressed tarnished silver setae that obscure sculpture. disk of clypeus sparsely covered with somewhat darker setae; terga I-II with brownish fasciae apically; vestiture of thorax (except on collar and propodeum posteriorly) and gaster brownish. Free margin of labrum slightly arcuate; clypeal lobe roundly trapezoidal (similar to fig. 41). its polished surface curving down to free margin, clypeal disk variably densely punctate. Frons dull. densely but shallowly punctate, punctures separated by about half a diameter; eye length greater than distance between eye notches (latter = 0.91 X eye length); UID 0.67X LID; ODD 0.42X HOD; flagellomere I length 2.5X apical width, remaining flagellomeres progressively shorter. Anterior margin of pronotum with broad, partly setose. polished lamella at level of pronotal lobe that narrows dorsad where it is paralleled behind by setose lamella that overhangs greatly elongate depression (similar to figs. 43. 45), length of depression about three-fourths width of collar, depression delimited by carina posteriorly; collar thicker than metanotum; scutum dull, with same dense punctation as frons; scutellum weakly shiny, somewhat less densely punctate than scutum; metanotum weakly shiny, more finely punctate than scutellum; propodeal dorsum with many oblique carinae basally that blend with many parallel, arcuate, cross-carinae along midline that change to dense, oblique striatopunctation laterally (similar to fig. 46), no median longitudinal carina; propodeal hindface with several coarse cross-ridges at petiole socket but striatopunctate above; propodeal side densely punctate, almost striatopunctate, punctures separated by half diameter or less, interspaces shiny; mesopleuron densely punctate, interspaces weakly shiny, punctures separated by one diameter or less and larger than elsewhere on body; metapleuron densely punctate, punctures smaller, shallower ventrad; plantulae present on tarsomeres II-IV. First recurrent vein interstitial between submarginal cells I-II, second recurrent ending on submarginal cell III. Tergum I more coarsely punctate than succeeding terga. punctures about same size as those on scutellum and separated by less than puncture diameter; posterior margin of tergum I broadly depressed (equal to width of fascia or breadth of metanotum at midline). Length 9 mm. Variation in females (9 specimens): Labrum sometimes truncate; UID 0.63-0.71 X LID; ODD 0.44X HOD in Ecuador specimen; propodeal dorsum with well formed median longitudinal carina in one specimen from Nova Teutonia, Brasil; cross-ridges of propodeal hindface present nearly to top in some specimens; apical depression of tergum r especially deep in Ecuador specimen; end point of recurrent veins highly variable with many combinations observed even in different wings of one specimen; length 7-9.5 mm. Male (I specimen): As in female except: clypeal lobe triangular; distance between eye notches 0.94X eye length; urD 0.86X LID; ODD 0.62X HOD; flagellomere r length slightly more than 2X apical width. III-V with vague sensory areas ventrally and slightly
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asymmetrically swollen; scutal punctures larger, deeper than those of frons; tergum VII narrowly truncate; genitalia as in figs. 50-52; apicoventral surface of gonostyle densely clothed with long, coarse, curved setae; penis valve slender, shorter than gonostyle. Discussion: The most diagnostic feature of fritzi is the anterolateral lamella on the pronotum (fig. 45). Pison nosferatu has only an anterodorsal setose lamella. The orientation of the carinae on the propodenl dorsum also differs between the two species. They are arcuately transverse in fritzi. but in nosferatu all the carinae are oriented obliquely at about a 450 angle (compare figs. 46-47 & 48-49). Range: Pison fritzi is known from Ecuador, southern Brasil and northwestern Argentina. ~:
Holotype female: BRASIL, Santa Catarina: Nova Teutonia, XII-l-1955, F. PIau mann (USNM). Para types (10 specimens): ECUADOR, Tungurahua: Pastaza Valley, 1600 m, VIII-25-87, 9, M. Cooper (COOPER). BRASIL, Santa Catarina: Nova Teutonia, various dates, 6 9, F. Plaumann (MCZ, UCD); Blumenau, 1-1885, 9, Hetschko (VIENNA); Guanabara: Represa do Rio Grande, VI-66, 9, M. Alvarenga (AEI). ARGENTINA, Salta: Rosario Lerma, II-85,
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than breadth of metanotum and less sharply defined. Length 9.5 mm. Variation in females (l specimen): 000 0.37X HOD; propodeal hindface with cross-carinae to top, upper carinae somewhat less coarse (fig. 48); length 9 mm. Male: --Unknown. Discussion: Pison nosferatu is similar to fritzi, but the absence of a polished lamella laterally on front edge of the pronotum is distinctive. The female clypeal lobe differs between the two species also (compare figs. 41-42). The punctation of nosferatu is somewhat coarser than that of fritzi, a difference difficult to appreciate without material of both species. The carinae of the propodeal dorsum are more oblique in nosferatu, being oriented at an angle of about 45 0 to the linear axis. In fritzi the carinae are more transverse (compare figs. 46-47 & 48-49). Range: Known only from Venezuela. ~:
Holotype female: VENEZUELA, Aragua: Rancho Grande, llOO m, 1-22/23-1978, blacklight in cloud forest, J. B. Heppner (USNM). One para type female with same data (USNM). Etymology: The name nosferatu, a noun in apposition, is from the 1922 German movie Nosferatu, based on the story of Dracula by Bram Stoker. In the movie, Nosferatu was a pseudonym for the vampire Dracula. Eremnon Group Pison eremnon Menke (Figs. 53-63) E'ison eremnon Menke, 1968a:5. Holotype female: Santarem, Brasil (CMP). Description: Body completely black, leading edge of fore and hindwings darkly infumate. Head and thorax (including the coxae, trochanters and femora) with much long, erect black setae. Tergum I and sterna with similar erect vestiture, but sparser on latter. Pale pubescence present only posterolaterally on propodeum and short. Head, thorax and terga I-II mostly bipunctate: large punctures scattered among many finer punctures (fig. 57). Nearly all punctures of disk of scutum and scutellum large (larger than any others on body), separated by one to six diameters, interspaces smooth, shiny. Propodeal dorsum covered with small punctures (no large punctures) that change to striatopunctation laterally. Terga III-V or VI densely, finely punctate, dull or weakly shiny. Tergum III with few slightly larger punctures scattered over surface. Sterna II-V shiny, irregularly punctate; punctures small, densest peripherally but mixture of large and small sparse punctures present on disk of female. Male sterna III-VI largely impunctate except laterally. Hindtibia with two to four short, very stout setae on outer surface that are borne on slight elevations, as well as several similar but more slender setae scattered over the surface in pits. Midtibia sometimes with single, stout seta.
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Head differs strongly between sexes: male with broad face and short clypeus in comparsion to female (see figs. 54-55). Median lobe of clypeus with broad, smooth shiny lip in both sexes that is not thickened. Female clypeus without brush. Eye asetose. Male UID O.72X LID, female UID 0.64X LID. DOD 3.2X HOD in male, 1.38X HOD in female. Occipital carina broadly incomplete ventrally. Female mandible with weak median tooth on the inner margin, but that of male edentate. Mandible socket nearly closed off by extension of hypostoma. Anterodorsal pronotal pit transversely elongate, deep, margined posterad by sharp lamelliform carina; width of pit about 0.66X UID. Collar as high as scutum, thicker than metanotum, in female with median elevation suggestive of Lyroda. Tegula impunctate on outer half. Propodeum with median longitudinal carina dorsally but no ridge present between spiracle and petiole socket. Propodeum posterolaterally with 5 or 6 coarse. transverse ridges above petiole socket that become progressively finer dorsad. Propleuron mostly punctate, but with small, shiny impunctate area present laterally on disk. Episternal sulcus curving forward ventrad attaining pleural margin. Hindcoxa with fine outer carina that evanesces basad. Tarsomeres II-IV with plantulae. Forewing marginal cell rounded distally (fig. 53). Submarginal cell I longer, narrower than typical for New World members of genus, resulting in petiole of submarginal cell II ending at midpoint of marginal cell (fig. 53). Recurrent veins terminate on submarginal cells II and III, respectively. Forewing media diverges from M+Cu basad of crossvein cu-a. Hamuli of hindwing not divided into two groups. Tergum I elongate, apical width slightly less than three-fourths its length, and hindmargin broadly depressed forming bandlike margin that is as broad as metanotum. Male abdomen unusual: tergum VI deeply emarginate, VII ending with pair of rounded lobes (fig. 56); sternum II strongly swollen in profile, it and subsequent sterna bearing subapical transverse setose flanges that diminish progressivly posterad, being represented on sterna IV-VI by setose depressions (figs. 58-59); sternum VIII ending in pair of blunt lobes (fig. 60). Genitalia laterally compressed; gonobase elongate, with longitudinal crest dorsally (figs. 61-62); gonostyle long, narrow, setose; and volsellar lobes elongate. setose apically, narrow in ventral view (fig. 63); aedeagus broad in dorsal view (fig. 61) but otherwise a fairly simple structure without defined head or ventral spine. Length 12.5-16 mm. Discussion: pison eremnon is the largest New World species of the genus, and it has many unique attributes. The shiny, irregularly bipunctate, black body, the dark erect setae of the head, thorax and legs, the wing venation and infumate leading edge of the forewing, the nearly closed mandible socket, the stout setae of the hindfemur, the elongate tergum I with its depressed bandlike hinpmargin, and the male abdominal features readily identify eremnon. The male is identified here for the first time and is represented by a single specimen from Pachitea, Peru (BERLIN). The long volsellar lobes are distinctive (fig. 63). The distally rounded forewing marginal cell and elongate submarginal cell I (fig. 53), the abdominal structure especially in the male (figs. 58-59), the nearly closed mandible socket, and head dimorphism (figs. 54-55) are unique attributes of eremnon that isolate it from all other the neotropical groups in the genus. I regard the nearly closed mandibular socket, the head dimorphism, and abdominal structures as apomorphies of the eremnon group. The nearly closed mandible socket is an interesting feature since it approaches the condition typical of the Philanthinae, where a closed socket is one of the prime characteristics that separate that subfamily from the Larrinae. The head of the male of eremnon is reminiscent of some species of Philanthus because of the notched eyes, the broad face and narrow clypeus. Strong head dimorphism occurs in the Old World species regale Smith and strandi Vasumatsu and their males have abdominal peculiarities suggestive of eremnon. But in regale and strandi the propodeum has a crenulate ridge between the spiracle and petiole socket, the mandible socket is broadly open. the mandible has a subbasal tooth on the cutting edge, the wing venation is different. and the male genitalia are
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dissimilar. A relationship between eremnon and these two Old World species, if any, does not appear close. Range: Colombia to Peru, Guyana and northern Brasil. Material examined (19 females, I male): COLOMBIA, Putumayo: Mocoa, 600 m, various dates, M. Cooper (BMNH); Villa Garzon, 8 km s Mocoa, various dates, M. Cooper (BMNH, COOPER); Amazonas: La Chorerra, VIII-l 4123-76, M. Cooper (BMNH); Leticia, VIII-21129-74, M. Cooper (BMNH); Meta: La Macarena, X-7/XI-29-76, M. Cooper (BMNH); Caqueta: Yuruyaco, 73 km sw F lorencia, II-5-79, M. Cooper (BMNH). ECUADOR, Morona-Santiago: Taisha, 500 m, 1-21-82, M. Cooper (COOPER). PERU, Pasco: Puerto Bermudez, 500 m, XI-8-84, M. Cooper (COOPER); Huanuco: Rio Pachitea, Tessman, (BERI_IN). GUYANA: Wanaina, N.W. District, III-31. J. Meyers (BMNH). Delicatum Group . Pison delicatum Menke, n. sp. - - (Figs. 64-76) Description, holotype male: Black, mandible yellowish brown, inner surface of foretibia brown; wings weakly stained but forewing with clear area near third submarginal cell. Body without long erect setae, and mostly covered by fairly dense, pale, short setae that impart a velvety appearance; clypeus and lower half of frons with dense, appressed silver setae that obscure sculpture; terga I-III with broadly interrupted silver fasciae. Labrum short, broad, free margin entire, slightly arcuate; clypeal free edge thickened, without prominent median lobe, but with very short, broad lobe defined by polished part of thickened edge, this lobe with weak, rounded, median tooth (similar to fig. 66); frons moderately swollen, dull, contiguously micropunctate, appearing granular; eye asetose, eye length 0.95X distance between eye notches; LID 0.83X UID; DOD 1.33X HOD; antenna clavate, flagellomere I length slightly more than 2X apical width (10:4), V-X broader than long; occipital carina incomplete ventrally, ending just before hypostomal carina. Pronotum with transversely elongate, polished anterodorsal depression whose length is equal to about half width of collar, this depression delimited anteriorly by high lamella (similar to fig. 67); collar slightly thicker than metanotum, noncarinate, but with shallow, anteromedian depression that narrows top of collar at midline; thorax shiny; scutal flange narrow, upturned; scutum densely micropunctate, punctures separated by about puncture diameter; tegula completely densely, micropunctate; scutellum margined anteriorly by narrow sulcus that is vaguely foveolate, punctures of scutellum sparser than on scutum; metanotum impunctate; propodeal dorsum with median longitudinal carina in shallow depression, surrounding surface smooth, sparsely covered with setose micropunctures; propodeal hindface with coarse cross-ridges that become weaker toward top; propodeal side with crenulate ridge between spiracle and petiole socket, side smooth, densely but very shallowly punctate, punctures about two diameters apart, lower end of propodeal spiracle surrounded by chain of small foveolae; propleuron finely punctate; episternal sulcus ending ventrally, not curving forward to anterior margin of mesopleuron; punctation of mesopleuron similar to scutum; mesopleural sulcus paralleled anteriorly by row of foveolae; metapleuron more sparsely, finely punctate than mesopleuron; metapleural flange not lamelliform. Forewing media diverging after crossvein cu-a; forewing with three submarginal cells, inner and outer vein lets of III narrowly separated on marginal cell, recurrent vein I ending on submarginal I. recurrent vein II ending on submarginal cell II. Legs smooth, devoid of conspicuous spiniform setae at apices of tarsomeres;
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tarsomere IV with tiny plantula; tarsal claw thick to just before apex where it terminates in a sharp, somewhat hooked point; midcoxa margined anterad by ridge associated with precoxal sulcus; dorsum of hindcoxa with strong inner carina, but outer carina reduced to short evanescent distal ridge. Distal edge of tergum I simple, surface of tergum I weakly shining, coriarius (see Harris, 1979) and obscurely punctate, punctures about a diameter apart; tergum II shiny, smooth, punctate; distal margin of tergum VII with broad. obtuse emargination; sterna III-V with discal mats of decumbent, long, white setae (as in figs. 68-69); sternum VIII with broad, apical V-notch (as in fig. 69); genitalia as in figures 73-76, aedeagus broad in dorsal view (fig. 76) and with unusual head (figs. 73-75) that has a nearly circumferential lamella ventrally that is joined at midventer by another lamella; gonostyle with usual long setae but subapically with area of dense, very short setae (fig. 74); volsellar lobes elongate, feebly setose. Length 5 mm. Variation in males (20 specimens): Mandible varies from yellow to nearly black. Mid and hindtibiae sometimes brownish within, sometimes legs completely black. Eye length sometimes as short as 0.90X distance between eye notches. LID sometimes only 0.76X UID. ODD 1.28-1.46X HOD. Ridges of propodeal hindface sometimes evanescent laterally and dorsad, occasionally hindface has only two or three ridges just above petiole socket with remainder smooth, punctate. Forewing media sometimes interstitial with cu-a. First recurrent vein sometimes interstitial between submarginal cells I-II in one or both wings, second recurrent rarely ending on third submarginal. Tergum I sometimes smooth, shiny, and clearly punctate. Length to 7 mm. Female (22 specimens): As in male except: edge of clypeus sometimes yellowish brown; vestiture sometimes dull silver; mandible with median cleft on inner margin, but no tooth; labrum arcuate; clypeus without prominent median lobe, free edge thickened at middle and produced into arcuate lower lip (figs. 64-65); eye length 0.92-0. 97X distance between eye notches; LID 0.74-0.84X UID; ODD l.24-1.53X HOD; flagellomere I length 2.5-3X apical width; punctation of scutellum about as dense as on scutum, metanotum with similar but sparser punctures; cross-ridges of propodeal hindface as variable as in male; vein lets of submarginal cell III sometimes meeting on marginal cell; first recurrent vein ending on first submarginal cell in 50% of the specimens, interstitial in about 50%, rarely both ending on second submarginal cell; length 6-8 mm. Discussion: Pison delicatum, a commonly collected species, has a distinctive velvety appearance due to a general covering of short setae on the body and an absence of erect setae. This look is enhanced by the dense micropunctures distributed generally over the body. Distinctive features of the species are the straight episternal sulcus, the ridge at the top of the propodeal side, the very elongate depression of the pronotum (fig. 67), and the hair mats on male sterna III-V (fig. 68-69). The elongate pronotal depression is similar to the condition in the fritzi group, and the lamella bordering it is probably homologous with the structure found in that group and the krombeini group. The lamellae of the aedeagal head are distinctive and make the genitalia unlike any other New World species (figs. 73-74). Apomorphies of the delicatum group are the broad face (fig. 64), the clavate antenna with distal flagellomeres wider than long, the non-spiny legs with peculiar broad tarsal claws (figs. 70-72), the straight (incomplete) episternal sulcus, the propodeal ridge, the long pronotal depression (fig. 67), and the hair mats of the male sterna (figs. 68-69). The antenna, absence of erect body setae, tarsal claws and completely punctate tegula are shared with the krombeini and stangei groups, but they have densely setose eyes (an apomorphy) and an episternal sulcus that curves forward ventrally to the anterior margin of the mesopleuron. The krombeini group differs from the delicatum group in possessing a broad lamella on the anterior
Menke: Pison in the New World
37
margin of the pronotum (an apomorphy), in lacking a lateral ridge on the propodeum, and in having a complete occipital carina. The male abdominal sterna have specialized areas in delicatum and krombeini groups, but they are dissimilar and presumably this is convergence. The delicatum and stangei groups share the propodeal ridge that extends between the propodeal spiracle and petiole socket, but in the stangei group there are only two submarginal cells and the eye is setose (both apomorphies). The pronotum has a simple, median pronotal pit anteriorly and the collar lacks an anteromedian depression in the stangei group. Pison agile and delicatum share a straight episternal sulcus, a completely punctate tegula, and have similar legs and antenna, but the setose eyes, lamelliform metapleural flange, and two submarginal cells are apomorphies of the agile group that separate it from the delicatum group. The occipital carina is a complete circle in the agile group. Ran~:
Pison delicatum occurs from Colombia and Venezuela to northern Argentina and southern Brasil. ~:
Holotype male: VENEZUELA, Zulia: Los Angeles del Tucuco, IV-1S/16-1981, A. S. Menke and L. Hollenberg (USNM). Paratypes (22 females, 20 males): COLOMBIA, YSlupes: Mitu, V-16-74, M. Cooper (BMNH); Meta: Cord. Macarena, I1-1S/28-76, M. Cooper (BMNH); Valle: Caula, VII-IO-7S, R. Wilkerson, malaise trap (CSOA), finca San Luis near Candelaria, IX-7/9-7S, J. Lattke (CAS). VENEZUELA, Borburata, Car., III-IS-40, P. Anduze (CU). ECUADOR, Napo: Coca on Rio Napo, V-6S, L. Pena (AMNH), Tena, XII-9/14-71, M. Cooper (BMNH), Limoncocha, VI-ISI28-76, S. & J. Peck (CNC); Zamora-Chinchipe: Timbara, IV-4-6S, L. Pena (AMNH). PERU, Cuzco: Quillabamba, XII-23/27-83, L. Huggert (PMA); Huanuco: Tingo Maria, VI-21-B2, malaise trap, Wasbauer and Sian sky (CSDA); Amazonas: Rodriquez de Mendoza, ISOO m, V-16-82, M. Cooper (COOPER); Junin: Valle Chanchamayo, 800 m., 1939, VIII-12-SI, W. Weyrauch (LILLO), San Ramon, Valle Chanchamayo, BOO m., I1-12-40, W. Weyrauch (AMNH). BOLIVIA, Beni: Rio Itenez about 4 km. above Costa Marques (Brasil), IX-12/IB-64, Bouseman and Lussenhop (AMNH); La Paz: Chulumani, 1700 m., IV-2-79, M. Cooper (BMNH); Coroico (BERLIN).~PARAGUAY: San Pedro Cororo-Rio Ypane, XII-1I4-83, malaise trap, M. Wasbauer (CSDA); Rio Ypane, Cororo, XI-79, M. Fritz (FRITZ). ARGENTINA, Misiones: El Dorado, XII-70, Foerster (FRITZ); Cataratas del Iguazu, XI-S/9-70, Porter and Stange, (LILLO); Salta: Oran, Abra Grande, X-18/2S-68, C. Porter (MCZ). BRASIL, Bahia: Itabuna, XI-1S-82, I1-21-83, F. P. Benton (BMNH); Sao Paulo: Ribeirao Pruto, X-IO-6S, R. Neilsen (UCD); Campinas, 1903, Hempel (VIENNA); Espirito Santo: Colatina, X-69, F. M. Oliveira (AE!). Non-type material (one headless specimen): BRASIL, Para: Almeirim, XII-16-02, A. Ducke (VIENNA). Etymology: The name delicatum is the Latin word meanings delicate, soft, or tender. Agile Group (Figs. 77, 81-8S) Description: Frons strongly swollen; face not broad, eye length slightly greater than distance between eye notches (fig. 77); labrum quadrangular, free margin entire; free margin of female clypeus with rounded median lobe whose edge is slightly thickened (fig. 77); female mandible with one inner tooth that is located slightly beyond midpoint; eye
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Contrib. Amer. Ent. Inst.. vol. 24. no. 3. 1988
densely covered with short setae; antenna clavate. outer flagellomeres broader than long; occipital carina a complete circle. narrowly separated from apex of hypostomal carina; pronotum with small. round anterodorsal pit but no lamella; scutal flange moderately broad; tegula completely punctate; scutellum bordered anterad by row of foveolae; propodeum punctate and with median longitudinal carina on dorsum but without lateral ridge at top of side; propleuron slightly gibbous posterolaterally. disk polished. impunctate; episternal sulcus straight. ending ventrad without curving forward; mesopleuron with subomaulus that often almost joins lower end of episternal sulcus; mesopleural sulcus paralleled anteriorly by row of foveolae; metapleural flange broadly lamelliform posteriorly (fig. 81); forewing media diverging after cu-a; forewing with two submarginal cells. second not petiolate. recurrent veins ending on submarginal cell I and II. respectively. or second interstitial between I-II; midcoxa bordered anterad by high ridge that is associated with precoxal sulcus; legs smooth. devoid of conspicuous spiniform setae at apices of tarsomeres; female fore trochanter without carina; hindcoxa with low inner carina dorsally. but without outer carina; hindtarsomere IV with very tiny plantula; tarsal claw thick to just before apex; tergum I simple apically; male sternum VIII narrow. entire or weakly indented at apex; genitalia laterally compressed. go no style simple but fringed ventrally with coarse. arcuate setae. volsellar lobes small. weakly setose. Included species: Pison agile in the North America. others (including agile) in the Oriental and Palearctic Regions: browni (Ashmead). differens Turner. hissaricum Gussakovskij. and possibly erythropus Kohl and rothneyi Cameron. Discussion: The following characterize the agile group. an Old World assemblage with one species adventive in North America: eyes setose. occipital carina complete. pronotum with only a small median pit anteriorly. subomaulus present. episternal sulcus straight. two submarginal cells. hindcoxal dorsum without an outer carina, and propodeum without ridge laterally. I have studied examples of browni and differens as well as agile. Although sternum VIII of the male is bluntly rounded with a slight indentation in agile (fig. 82), it can be narrower and more pointed as in browni from the Philippine Islands (see fig. 239 in Tsuneki, 1983). Apomorphies of the agile group are the setose eyes, the clavate antenna. the two submarginal cells. the presence of a subomaulus. the straight episternal sulcus. the broad metapleural flange (fig. 81). and the form of the tarsal claws. The subomaulus is merely a short spur in browni. Earlier I (Menke, 1968b) called this the koreense group but Krombein (1979) indicated that agile was a senior synonym of Radoszkowski's name. Pison agile (Smith) (Figs. 77, 81-85) Parapison agilis Smith, 1869:300. Holotype female: Ceylon (BMNH). Paraceramius koreensis Radoszkowski. 1887:433. Holotype female: Koree (Mus. Krakow?) Pison koreense. Krombein. 1958a: 166. Adventive in North America (Illinois, Virginia). Pison koreense, Krombein. 1958b: 189. Catalog. PB.raCeramius koreensis. Richards. 1962:118. Paraceramius preoccupied. Krombeinielleum proposed as replacement name. Pison koreensis, Iwata. 1964: 1. Biology. Pison koreense. Krombein. 1967:394. Records from Maryland and Michigan. Pison koreense. Menke. 1968a:3. 7. Taxonomy. listed. Pison koreense. Menke. 1968b:llOO. 1102. Taxonomy, keyed. Pison koreense. Sheldon, 1968:107. Biology. Pison koreense. Bohart and Menke. 1976:333. 337. Listed.
Menke: Pison in the New World
39
Pison agile. Krombein. 1979:1641. Synonymized koreense with agile. record from Kansas. Discussion: The following combination of characters separate agile from all other New World Pison: eye densely setose. metapleural flange broadly lamelliform (fig. 81). forewing with only two submarginal cells. propodeum without a dorsolateral ridge. and surface of tergum I dull in contrast to the shiny surface of II. Sternum VIII and the male genitalia are shown in figures 82-85. This is the only Pison known in North America, and it is adventive from the Oriental Region (Krombein. 1958a). I have not studied the types of koreense or agile. Biology: Iwata (1964) reported on agile nests in Japan. and Sheldon (1968) observed the species in Illinois and described the biology in detail. including illustrations of the larva. Both authors used the name koreense. Iwata illustrated the cells of agile. and described them as "very thin and fragile" and said that "clearly [they] may be the ..... most fragile mud cell[s] made by Japanese Aculeata." He said that a nest (16 studied) consisted of one to 21 separate but contiguous cells arranged irregularly (average number of cells/nest = 7.6). They were glued to vertical protected surfaces. and the exterior surface had a "ripple relief." Iwata found 6 immature spiders of the genus Araneus in one cell. Sheldon reported that agile made its nests in sheltered situations also. and even used empty cells of Trypoxylon politum (Say) nests. One to 12 fragile cells. arranged randomly. made up a nest. Adult spiders of the genus Dictyna representing two species. were the prey found in four cells. but immatures of the same genus were also provisioned. The average number of spiders per cell was 25.2. Sheldon stated that the female carried the paralyzed spider in her mandibles. The wasp egg was laid on the last spider provisioned. The eulophid parasitoid Melittobia chalybii Ashmead was reared from one cocoon of agile that Sheldon found in an unsealed cell of Trypoxylon politum (Iwata found no parasitoids in his observations). Range: Apparently widespread in the Oriental Region (Sri Lanka. India) and eastern Asia (China. Japan. Korea). In North America agile is known from Kansas. Illinois. Michigan. New York. Maryland. and Virginia. Material examined (12 specimens): ILLINOIS: Palisades Park (USNM); Kickapoo State Park (USNM). MICHIGAN: Gun Lake Mal. Tr. (BMNH). NEW YORK: Ithaca (CU); Lewisboro (AMNH). VIRGINIA: McLean (USNM. BMNH). Stangei Group (Figs. 9. 12.78.86-108) Description: Scape. mandible. mouthparts. and most of fore and mid legs yellowish brown. Body devoid of erect setae; frons not swollen; face not broad. eye length greater than distance between eye notches; free margin of labrum arcuate or quadrangular; female clypeus not double-edged but margin sometimes thickened. a median lobe weakly or not differentiated; male clypeus tridentate. thickened; female mandible with one inner subapical or median tooth or simple; eye densely covered with short setae; antenna clavate. outer flagellomeres broader than long; occipital carina usually incomplete below. ending just before reaching midventral line. occasionally a complete circle narrowly separated from apex of hypostomal carina; pronotum with small round or transversely elongate pit anterodorsally; pronotal collar without anteromedian depression but sometimes with obliquely transverse ridge; scutal flange
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narrow, upturned; parapsidal lines sulci form in female; tegula completely micropunctate; scutellum margined anteriorly by row of contiguous foveolae; propodeal side delimited dorsad by crenulate ridge that extends from petiole socket area to spiracle; propleuron punctate, with gibbosity posterolaterally; episternal sulcus curving forward ventrally, attaining anterior margin of mesopleuron but often weakly impressed there; mesopleural sulcus paralleled anteriorly by row of foveolae; metapleural flange narrow; forewing media diverging after cu-a; forewing with two submarginal cells (three in rare exceptions, figs. 87-88), II not petiolate in two-celled wing (figs. 12, 86); both recurrent veins ending on submarginal cell I, or second interstitial between I-II, or ending on II; midcoxal cavity bordered anterad by high ridge associated with precoxal sulcus; legs smooth, devoid of conspicuous spiniform setae at apices of tarsomeres; tarsomeres apparently without plantulae; tarsal claw thick to just before apex; female foretrochanter not carinate ventrally; dorsum of hindcoxa with low or very weak inner carina and no outer carina or only an apical vestige; apical rim of tergum I simple; male sternum VIII narrow, apex truncate or with shallow emargination; gonostyle of male genitalia long, slender, simple, setose ventrally, volsellar lobes small, aedeagus without ventral spine. Included species: Pison abathes, duckei, plaumanni and stangeL Discussion: The following characterize the stangei group: eye densely setose (fig. 9), pronotum with small median pit anteriorly (fig. 95), episternal sulcus complete, forewing usually with only two submarginal cells (but see below), and propodeum with dorsolateral ridge. In this assemblage the true second submarginal cell has been reduced to the point of disappearing, but in one male a tiny second submarginal is present in both wings (figs. 87-88). The eye notch is unusually shallow in one species, abathes. Members of the stangei group are among the smallest species of Eison known. Apomorphies of the group include the setose eyes, the clavate antenna, the form of the tarsal claws, the two submarginal cells, and lateral propodeal ridge. In terms of species discrimination, the stangei group is the one of the most vexing assemblages in the New World fauna. More material has been available than when I first treated the group (Menke, 1968b), but differences between stangei and duckei are still not as precise as one would like, and the male of the latter is still unknown. Pison plaumanni remains the most distinctive species and its male is now known. One new species has been discovered. Finally a male from Ecuador (AMNH) has three submarginal cells, the second represented in each forewing by a much reduced cell (figs. 87-88). The wing condition of this specimen may be a freak occurrence, but in any case it is probably an undescribed species. The male genitalia are fairly similar in pIau manni, stangei and the specimen with three submarginal cells. Pison plaumanni Menke (Figs. 9, 12,89-90,95,101-104) Pison plaumanni Menke, 1968b: 1105. Holotype female: Nova Teutonia, Santa Catarina, Brasil (UCD). Description: Pedicel, basal part of flagellum, and clypeal margin often yellowish brown. Female clypeus with weakly defined lobe: it is thickened, truncate, and its edge has a transverse dimple (fig. 89; the clypeal outline in Menke, 1968b, fig. 7, is inaccurate). Male clypeus as in fig. 90. UID 0.68-0.69X distance between the eye notches in females, O.72X in males. LID 0.69-0.71 X UID in females, 0.63-0.66X in males. Ocellocular distance greater than other species of stangei group (ODD 1.38-1.65X HOD). Occipital carina broadly interrupted ventrad in comparison with other species, gap equal to length of foretarsomere I or only slightly shorter (17:20). Anterodorsal
Menke: Pison in the New World
41
pronotal pit circular to transverse, usually margined posterad by carina. Pronotal collar rounded laterally, no trace of transverse ridge or humeral angle (fig. 95). Disk of propodeal dorsum smooth, punctate, with median longitudinal carina that is often in depression; carina usually reaches propodeal apex or nearly so, and frequently many short cross-carinae associated with it. Propodeal hindface varies from punctate to transversely ridged and punctate. Propodeal side smooth, punctate, without ridges except sometimes just beneath spiracle. Second recurrent vein interstitial between submarginal cells I-II, or ending just inside l. Male sternum VIII slightly emarginate at apex, lateral margins parallel or slightly diverging basad (fig. 104). Male genitalia similar to §.tangei but aedeagus without midventral ridge (fig. 102). Length 5-6.5 mm. Discussion: This is the largest species in the stangei group. The broad ocellocular distance, the rounded humerus of the pronotal collar (and absence of an oblique ridge), and smooth propodeal dorsum and side are the most diagnostic features. The eighth sternum of the male is not constricted before the apex in contrast to that of stangei. I have examined two females from Mocoa, Putumayo, Colombia (BMNH) that resemble plaumanni in the clypeus, the broadly interrupted occipital carina, the rounded humerus of the pronotal collar, the propodeal sculpture, and body size. However, the ocellocular distance is only slightly greater than an ocellus diameter (ODD 1.1l-1.l4X HOD), the LID is O.76-0.78X the UID, and the UID is only 0.61-0.62X the distance between the eye notches. Thus these specimens may represent an undescribed species. Range: Pison plaumanni is known only from eastern and southeastern Brasil. Material examined (26 females, 12 males): BRASIL, Bahia: Itabuna, VIII-7-83, III-18-84, F. Benton (BMNH); Aguas Vermelhas, XII-83, M. Alvarenga (AE!). Sao Paulo: Sao Paulo, various dates, V. Alin (MLSU). Santa Catarina: Nova Teutonia, various dates, F. Plaumann (UCD, MCZ, USNM, BMNH). "Brazilia", 1886 (USNM). Pison stangei Menke (F igs. 86, 91-92, 94, 96, 105-108) Pison stangei Menke, 1968b:ll02. Holotype male: Amaicha, Tucuman, Argentina (LILLO). Description: Antenna and fore and midlegs occasionally entirely or largely black. Female clypeus with prominent median lobe the edge of which is variably thickened (fig. 91). Male clypeus as in fig. 92. UID 0.59-0.62X the distance between eye notches in females, 0.62-0.64 in males. LID 0.74-0.81X UID in females, 0.76-0.79X in males. ODD 1.08-1.27X HOD. Occipital carina often a complete circle narrowly separated from apex of hypostomal carina, but sometimes weakly complete or narrowly incomplete at midventral line. Anterodorsal pronotal pit circular, not margined posterad by carina. Pronotal collar rounded laterally although weak transverse ridge present (figs. 94, 96). Propodeal dorsum with median carina that fades posterad, usually ending well before apex; disk of dorsum variably sculptured: coarsely to finely obliquely ridged or striatopunctate, or ridging becoming transverse on apical half to two-thirds, or surface obscurely striatopunctate, or even mostly just punctate. Propodeal hindface transversely ridged, ridges sometimes evanescent laterad. Propodeal side obliquely or horizontally ridged above, punctate below. Second recurrent vein ending on submarginal cell I or sometimes interstitial between I-II. Male sternum VIII slightly concave at apex, lateral margins slightly constricted
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subapically (fig. 108). Male genitalia as in figs. lOS-1 07, aedeagus with midventral ridge (figs. lOS-106). Length 4.S-6 mm. Discussion: The rounded humerus of the pronotal collar and the fine ridges on the upper half of the propodeal side are the most obvious diagnostic features of stangeL The weak transverse ridge on the collar (fig. 96) is a condition intermediate between QJilumanni, which has none (fig. 9S), and duckei and abathes, both of which have a well developed oblique ridge (figs. 97, 99). Except for the difference in the collar. stangei is similar to duckei, and their head measurements are essentially identical. The occipital carina is often a complete circle in stangei. but it is always interrupted in duckei. When I described stangei (Menke, 1968b), I used the ridging of the propodeal dorsum as a diagnostic feature for separation from duckei, the latter having a mostly punctate dorsum. But the sculpture of the dorsum varies considerably in stangei, some specimens being mostly punctate and therefore similar to duckei. Thus propodeal sculpture is not reliable for separating these two species. Range: Pison stangei is known only from northwestern Argentina and Bolivia. Material examined (14 females, 4 males): BOLIVIA, Tarija?: Las Carreras, I-4-S6 (KU). ARGENTINA, Salta: Rosario de Lerma, XI-10/18-83, M. Wasbauer (CSDA), Pocitos, 1-1971. M. Fritz, Martinez and Fritz (USNM). Tucuman: Horco Molle, Parque Sierra San Javier, 700 m, various dates, L. A. Stange (ULLO), Jardin del Ullo, IX-19-67, C. Porter (MCZ), Tucuman, XI-24-6S, L. A. Stange (LILLO); Santiago del Estero: Suncho Corral, XII-28-7S, L. Stange (LILLO). Pison duckei Menke ----cFigs. 97-98) Pison duckei Menke, 1968b: 1103. Holotype female: Para, Brasil (VIENNA). Description (female only): Pedicel and flagellomere I usually yellowish, clypeal margin yellowish in type, hindleg largely yellowish brown in material from Trinidad, midleg black above in specimen from Brasilia. Clypeus with prominent median lobe whose edge is thickened (similar to ~tangei). UID 0.60-0.66X distance between eye notches. LID 0.70-0. 79X UID. ODD 1.1-1.31 X HOD. Occipital carina interrupted ventrad. Anterodorsal pronotal pit circular, margined posterad by short carina. Pro notal collar with oblique, transverse ridge that makes humerus angulate when viewed from above (figs. 97 -98). Propodeal dorsum punctate, with median longitudinal carina that ends at middle or slightly beyond, many short cross-carinulae associated with median carina, base of dorsum with up to 10 oblique carinae that only rarely extend onto disk (one Trinidad specimen); hindface with fine cross-carinulae that become evanescent dorsad; propodeal side variable: sometimes with several horizontal ridges only beneath spiracle, sometimes upper area more extensively ridged. Both recurrent veins ending on first submarginal cell. Length 4-4.S mm. Discussion: Pison duckei is similar to stangei. The essential features of duckei for separation from stangei are: pronotal collar with obvious oblique transverse ridge that makes humerus clearly angulate (fig. 97), occipital carina interrupted, and propodeal dorsum mostly punctate. The degree of the pronotal angulation varies however, and the punctate propodeal dorsum is approached by some specimens of
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stangeL The sculpture of the propodeal side is often similar between the two species. The male of duckei is unknown. The male with three submarginal cells discussed under the stangei species group is not the unknown male of duckei, although its head measurements fall within the range of the species. The sculpture of its propodeal dorsum is more like stangei, the humerus of the pronotal collar is not angulate, and of course the wing has three submarginal cells. Range: Pison duckei is known from scattered localities between Panama and central Brasil. Material examined (11 females): PANAMA: Las Cumbres, XII-BI2B-B2, H. Wolda (PMA). COLOMBIA, Tolima: Armero. I-30/II-5-77. E. L. Peyton (USNM). VENEZUELA, Guarico: Hato Masaguaral. 44 km s Calabozo, 5-3/1O-B5, Menke & Carpenter (USNM). TRINIDAD: st. Augustine, various dates, E. Callan, J. Noyes (BMNH). BRASIL, Para: Para (Belem). I-IS-Ol, A Ducke (holotype) (VIENNA); Federal District: Brasilia, Lago Sui, 1II-7-77 , A. Raw (BMNH). Pison abathes Menke. new species (Figs. 7B, 93. 99-100) Description, holotype female: Black, following yellow brown: palpi, mandible. labrum, clypeal margin, scape and pedicel beneath, margin of pronotal lobe, fore and midlegs except coxae, and hind trochanter . Labrum trapezoidal in shape, shallowly concave (fig. 93); clypeus with arcuate median lobe whose margin is slightly thickened; eye notches shallow (fig. 7B). UID O.BX distance between eye notches; LID 0.6SX UID; ODD 1.27X HOD; occipital carina complete but weak opposite apex of hypostomal carina and separated from latter; anterodorsal pit of pronotum circular, smaller than hindocellus; collar with oblique, transverse ridge that makes humerus angulate in dorsal view (figs. 99-100); scutum, scutellum and metanotum weakly shining, finely, nearly contiguously punctate. punctures largest on scutum; propodeal dorsum shiny, with about B short, coarse carinae at base, middle one extending only about one-third distance to apex. disk of dorsum irregularly transversely carinulate changing to punctation laterally, apex of dorsum delimited from hind face by short carina; hindface shiny, with coarse . cross-ridges; propodeal side longitudinally carinulate on upper one-fifth, remainder punctate; both recurrent veins ending on submarginal cell I. Length 4.5 mm. Variation in females (3 specimens): Clypeus completely black (Bolivia), hindfemur yellow brown (Bolivia, Guyana). Edge of clypeal lobe not thickened (Bolivia, Guyana). UID 0.76-0. 7BX distance between eye notches; LID 0.67-0.70X UID. ODD 1.13-1.27X HOD. Propodeal dorsum largely punctate in Bolivian specimen. median carina extending half distance to apex, the latter without transverse carina; dorsum in Guyana specimen more like holotype but median carina almost reaching apex, the latter with only suggestion of a transverse carina. Propodeal side with only two carinulae just beneath spiracle (Bolivia, Guyana). Second recurrent vein interstitial in one wing of Guyana paratype. Length 4-4.5 mm. Male: Unknown.
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Discussion: The shallow eye notches, angulate pronotal humerus, and trapezoidal labrum are the salient features of abathes. The variation seen in the sculpture of the propodeal dorsum is disconcerting and it may be that the Bolivia and Guyana material is not con specific with the Ecuadorian females. However, there is also considerable variation in the propodeal sculpture of stangei. It is possible also that the shallow eye notches of abathes simply represent extreme variation of duckei, the only other species with angular pronotal humeri. But lacking intergrades I have decided to treat these specimens as a new species. Range: Ecuador, Bolivia and Guyana. ~:
Holotype female: ECUADOR, Pichincha: Rio Palanque Res. Sta., Il-1983, malaise trap, Sharkey and Masner (PMA). Paratypes (three females): ECUADOR, same data as type (PMA). BOLIVIA, Santa Cruz: Rio Mamore, 2 km n mouth of Rio Chapare, VII-31-65, J. Bouseman (AMNH). GUYANA: Mazaruni. clearing, in colony house, VIII-26-37, o. Richards (BMNH). Etymology: The name abathes is a Greek word that means shallow, a reference to the distinctive eye notches of this species. Krombeini Group (Figs. 79-80, 109-116) Description: Frons not strongly swollen; face broad, eye length about 10% shorter than distance between eye notches (figs. 79-80); labrum transverse, hidden, free margin arcuate or obtusely triangular, entire; free margin of female clypeus not thickened or double-edged, outline a low obtuse triangle or with median lobe; female mandible with subapical inner tooth; occipital carina a complete circle, broadly contiguous with apex of hypostomal carina; eye densely covered with short setae; antenna clavate, outer f1agellomeres broader than long; anterior edge of pronotum with broad, thin, setose lamella that extends laterad to level of pronotal lobe; pronotum without anterodorsal pit; scutal flange broad, upturned; tegula evenly punctate; scutellum with unpitted narrow sulcus anteriorly; propodeum punctate, with median longitudinal carina on dorsum but without lateral ridge, hindface without cross-ridges; propleuron with short, high lamelliform projection posterolaterally; episternal sulcus curving forward ventrally, attaining anterior margin of mesopleuron; mesopleural sulcus paralleled by series of foveolae; forewing media diverging after cu-a; forewing with three submarginal cells, II receiving both recurrent veins; midcoxal cavity delimited anterad by high ridge associated with precoxal sulcus; legs smooth, devoid of conspicuous spiniform setae at apices of tarsomeres; female fore trochanter with carina ventrally; outer dorsal carina of hindcoxa represented by high apical lamella; tarsi without plantulae; tarsal claw thick to just before apex; tergum I with reflexed apical rim that is lamelliform; male sternum VIII semicircularly emarginate apically, flanked by two prongs (fig. 110); male genitalia as in figs. 113-116, gonostyle setose, with narrow, ventral, SUbapical lobe (fig. 115); volsellar lobes large, setose apically; aedeagal head with 'ventral process (see fig. 13 in Menke, 1968b). Included species: Pison krombeini and neotropicum.
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Discussion: The following define the krombeini group: face broad, eye setose, occipital carina complete, pronotum with setose lamella along anterior margin, propodeum without ridge laterally, forewing with three submarginal cells, tegula entirely punctate, tergum I apex reflexed, and male sternum VIII emarginate, bounded by prongs. The tergal character needs some elaboration. The lamella-like apical margin of tergum I is not the real rim. The true edge of the tergum can be seen beneath this broad, upturned flange - thus, tergum I is really double-edged. Earlier I (Menke. 1968b) described the female mandible as having two subapical teeth, but there is only one tooth in krombeini, and only a suggestion of a second subapical tooth in neotropicum. Apomorphies of the group are the broad face. the setose eye, the pronotal lamella, the carinate female foretrochanter, the reflexed and double-edged margin of tergum I, and the two prongs at the apex of male sternum VIII. A similar but asetose pronotal lamella also occurs in the pilosum group. The male of krombeini has transverse depressions on sterna III-IV, but the male of neotropicum is unknown; thus, at present, it is impossible to list this specialization as an additional apomorphy for the group. Because I described both species earlier (Menke, 1968b), I am presenting only brief diagnoses and distribution records here. Pison krombeini Menke (Figs. 79, 109-llO, 112-116) Pison krombeini Menke, 1968b: 1105. Holotype male: British Honduras (= Belize) --(USNM). Diagnosis: The dense metallic setae of the propodeal dorsum and hindface is the most obvious recognition feature. The female clypeus (fig. 109) differs from that of neotropicum (fig. lll). The thorax is shiny, the pronotal collar has a prominent ridge anterolaterally, and the parapsidal and admedian lines of the scutum are deeply impressed. The metapleural flange is narrow. Terga I-II have broad silver or golden apical fasciae, and sternum I has appressed silver setae that are directed basad. The forewing has a cloud at the apex of the first submarginal cell and the leading edge of the wing is faintly infumate. The inner and outer vein lets of submarginal cell III are separated on the marginal cell by an ocellus diameter or less, sometimes joining there, or even forming a petiole. The depressions on male sterna III-IV are presumably diagnostic also (fig. 112). The aedeagus has a large ventral prong (not visible in figs.), the apices of the two penis valves curl away from each other (figs. 114-115), and the gonostyle has a curled fingerlike process on its inner, ventral margin near the apex (figs. 114-115). Length 6-7.5 mm. Range: Southern Mexico to Colombia. Material examined (13 females, 3 males): MEXICO, Vera Cruz: Cordoba (GENEVA); Tabasco: Teapa, March, H. H. Smith (BMNH); Yucatan: Chichen Itza, VII-20-52, J. & D. Pallister (AMNH). BELIZE: "British Honduras", 1-08 (USNM). GUATEMALA: El SaIto, Escuintla, VI-28-34, F. X. Williams (BPBM), Helvetia, San Sebastian, IV-21122-31, J. Bequaert ((MCZ), Sta. Emilia, Pochuta, IIIIII-3I, J. Bequaert (MCZ). COST A RICA, Santa Rosa Park, II-13-78, D. Janzen (AEI). PANAMA: Barro Colorado I., VIII-15/IX-26-28, P. Rau (note 7756) (USNM). COLOMBIA, Arauca: Tame, VII-8/17-76. M. Cooper (BMNH); Magdalena: Rio Frio, VII-13-27, G. Salt (BMNH); Valle: Cali and vic., 3-4000 feet, 1-8/20-72, H. Evans (MCZ).
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Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988 Pison neotropicum Menke (Figs. 80, Ill)
Pison neotropicum Menke, 1968b: II 06. Holotype female: Rio de Janeiro, Brasil (USNM). Diagnosis (male unknown): Distinctive features of neotropicum are the shape of the female clypeus (fig. Ill), the dull thoracic dorsum, absence of appressed metallic setae on the propodeal hindface and apex of the dorsum, absence of silver setae on sternum I and absence of metallic fasciae on terga I-II. The collar has a transverse ridge anteriorly, but it is not prominent laterally. The admedian and parapsidal lines of the scutum are not depressed. The metapleural flange is about as broad as an ocellus diameter. The leading edge of the forewing is moderately infumate, and the inner and outer veinlets of submarginal cell III are widely separated on the marginal cell. Length 8.5 mm. No additional material has been found since the species was described. Range: Known only from southeastern Brasil. Material examined (6 females): BRASIL, Guanabara: Rio de Janeiro, X-38/I-39, R. C. Shannon (USNM, UCD). Euryops Group (Figs. II, 117-130) Description: Frons not swollen; face broad, eye length 10-20% shorter than distance between eye notches (fig. 117, 119); female clypeus with roundly truncate median lobe, its apex with lower lip of variable shape; male clypeus with sharp, triangular median lobe; female mandible with cleft near middle of inner margin, but no tooth; labrum with deep, U-shaped emargination resulting in two apical, setose, fingerlike lobes; occipital carina joining apex of hypostomal carina; pronotum without anterodorsal lamella or pit; lateral margin of scutum a broad upturned flange; tegula uniformly punctate; scutal flange broad; propodeal dorsum without median longitudinal carina; propodeal side with crenulate ridge that extends from petiole socket nearly to spiracle; disk of propleuron largely impunctate, shiny; lower end of episternal sulcus curving forward ventrally, usually attaining anterior margin of mesopleuron; dorsum of hindcoxa with strong inner carina, but outer carina reduced to short evanescent distal ridge; forewing media diverging from M+Cu after crossvein cu-a although sometimes nearly interstitial; forewing usually with three submarginal cells, rarely two (fig. II); tergum I narrowly double-edged apically; male sternum VIII with broad, shallow emargination apically; male genitalia laterally compressed, aedeagus with broad, apically rounded, laterally compressed head whose ventral margin has a hooklike angle (not visible in SEM photo, figs. 128-130), volsellar lobes slender, acuminate, apical half of gonostyle narrow, setose ventrally (figs. 128-129). Included species: Pison eu, euryops, lillo, and styphopteron. Discussion: The following define the euryops group, one of the most distinctive in the New World fauna: face broad, labrum with U-shaped emargination, pronotum without anterodorsal pit, propleuron largely impunctate, tegula completely punctate, and tergum I double-edged. The broad face, labral form and double-edged tergum are apomorphies. The labral and tergal character states also occur in the pilosum group but neither is universal there. Members of the euryops group have very similar sculpture and vestiture. Species
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differences are found mainly in clypeal shape, facial proportions, flagellar features, and sculpture of the propodeal hindface. The loss of the second submarginal cell in one species, styphopteron, appears to be a consistant feature, but only three specimens are known. The genitalia display relatively minor differences between the four species, but insufficient male material of lillo, styphopteron and eu has The hampered assessment of their usefulness in species discrimination. asymmetrically swollen flagellomeres in males of euryops are distinctive, but the few males available of the other species does not permit positive conclusions on the reliability of the apparent antennal differences observed. Pison eu Menke, n. sp. (Figs. 117-11B, 121, 12B-130) Description, holotype female: Black; wings clear except for cloud over submarginal cell II. Clypeus and lower half of frons with dense, appressed, silver setae that obscure sculpture; terga I-II with silver apical fasciae that are interrupted at middle; erect setae of body pale, longest on metanotum and propodeum. Clypeal lobe arcuate, its lower lip with broad, shallow emargination (similar to fig. lIB). Frons shiny , finely, densely punctate (punctures 1 diameter apart or less); eye length 0.8BX distance between eye notches; UID O.77X LID. ODD 0.B5X HOD. Flagellomere I length 3.SX apical width, remaining flagellomeres progressively shorter. Thorax shiny, scutum and scutellum very finely punctate, punctures sparser than those of frons (1-4 diameters apart); metanotum impunctate; propodeal dorsum largely smooth but disk with sparse, pinprick punctures that become denser peripherally and along midline; propodeal hindface with several transverse ridges just above petiole socket, upper part of hindface smooth, with scattered pinprick punctures; propodeal side smooth, uniformly covered with pinprick punctures (several diameters apart). Mesopleuron smooth, densely punctate, punctures 1-2 diameters apart anteriorly and about same size as those on scutum, becoming sparser, finer toward mesopleural sulcus; row of foveolae along lower half of mesopleural sulcus. Metapleuron same as propodeal side. Forewing with three submarginal cells, recurrent vein I ending on submarginal cell II, recurrent vein II ending on submarginal cell III (nearly interstitial in right wing). Tergum I with fine pinprick punctures similar to those on propodeal dorsum; II more densely punctate. Length 7.5 mm. Variation in females (15 specimens): Lower clypeal lip not indented in single specimen from Oaxaca. UID 0.76-0.BOX LID. Punctation varies in coarseness from specimen to specimen. Tergum III sometimes has apical fascia. Recurrent vein II is interstitial between submarginal cells I! and III in eleven specimens. Occasionally forewing uniformly but weakly infumate. Length up to 9.5 mm. Male (2 specimens): As in female except: mandible as in fig. 121; clypeal lobe as in fig. 121. UID 0.93-0.95X LID; ODD = HOD in Peruvian specimen. Flagellomeres VI-VI! each with weak linear tylus (Venezuelan specimen) or narrowly linear shiny area (Peruvian specimen). Genitalia as in figs. 12B-130. Length 6.5 mm. Discussion: The female clypeal lobe and the shape of its lower lip are immediately piagnostic in that sex (fig. liB). The non-ridged upper part of the propodeal hindface is also distinctive in both sexes, although styphopteron is similar. The latter species,
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Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988
however, has only two submarginal cells and other differences. The upper interocular distance is 0.73-0.80X the lower interocular distance in females of styphopteron and eu. In euryops the UID and LID are essentially equal. Flagellomere I is more than 3X as long as wide in females of eu and usually slightly less than 3X as long as wide in euryops. This flagellomere is about 4X as long as wide in females of styphopteron. The difference in the flagellum of the two males is disconcerting, but no other differences between them are apparent, and I am assuming that they are con specific. Range: Pison eu occurs from southern Mexico to southern Peru and eastward to Suriname, but I have no records from the Amazon basin proper. ~:
Holotype female: COLOMBIA, Narino: Barbacoas, IV-2-1974, M. Cooper (BMNH). Paratypes (17 specimens): MEXICO, Oaxaca, 44 mi. w Tehuantepec, VII-21-52, 'i', Gilbert and MacNeil (UCD). HONDURAS: La Ceiba, 'i', F. J. Dyer (USNM). COSTA RICA, Guanacaste: Santa Rosa Park, VII-15-77, 'i', D. H. Janzen (AEI); La Selva Res. Sta., VI-1l1l7-86, 'i', Hanson and Bohart (USU). COLOMBIA, Meta: Restrepo, VI-18-74, 'i', L. A. Stange (LILLO); Rio Duda, III-8/12-76, 'i', M. Cooper (BMNH); Arauca: Tame, VII-II7-76, 'i', M. Cooper (BMNH); Putumayo: Villa Garzon, 400-550 m, XII-25-87, 'i', M. Cooper (COOPER). PERU, Huanuco: Tingo Maria, VI-23-82, C!, malaise trap, Wasbauer and Siansky (CSDA); Cuzco: Quincemil, 750 m, X-16/31-62. 'i', L. Pen a (MCZ). VENEZUELA, Aragua: El Lim6n, 480 m, IV-24129-V-I-73, C!, 2'i'. malaise trap, C. Rosales (UCM). GUY ANA: Essequibo River, Moraballi Creek, X-8-29, 2'i' (BMNH). SURINAME: Paramaribo, VIl-20-60, XI-22-57, 2'i'. P. v. Doesburg (LEIDEN). Etymology: The name eu, treated as a noun in apposition, is based on the Greek prefix meaning true or good, a reference to the fact that the species is valid. Pison euryops Menke, n. sp. (Figs. 119-120, 122. 126-127) Description, holotype female: Same as described for eu except: forewing lightly infumate except for clear spot at submarginal cell III; tergaI-II with broadly interrupted fascia; clypeal lobe roundly trapeziform, its lower lip arcuate (similar to fig. 120); eye length 0.80X distance between eye notches; UID essentially equal to LID; OOD 1.31 X HOD; flagellomere I length less than 3X apical width (19.5:7.5); punctures of scutum and scutellum I to 2 diameters apart, metanotum with sparse pinprick punctures; propodeal dorsum punctures slightly larger than those on scutum, 1 to 2 diameters apart; propodeal hindface completely covered by coarse, transverse ridges; second recurrent vein interstitial between submarginal cells II and III; tergum r with punctation similar to scutum. Length 8 mm. Variation in females (15 specimens): Eye length up to 0.87X distance between eye notches. OOD usually 1.25X HOD, but OOD = HOD in one specimen from Guanabara. Thoracic punctation varies in degree of coarseness from specimen to specimen. Recurrent vein I occasionally interstitial between submarginal cells I-II. Length 6.5-9 mm. Male (9 specimens): As in female except: inner margin of mandible sinuate (fig. 122); clypeal lobe a simple triangle, sides straight (fig. 122); LID 0.83-0.95X UrD; OOD 1.33X HOD or slightly more; flagellomeres III-VI with linear. asetose, shiny areas ventrally (fig.
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127), IV-VII or VIII swollen toward apex in profile, V-VI most strongly so (fig. 126); second submarginal cell greatly reduced in two specimens from Argentina (smaller than ocellus, absent in one wing of one specimen); first recurrent vein ending most commonly on submarginal cell II, but sometimes interstitial or ending on I. second recurrent vein usually interstitial but sometimes ending on submarginal II or III; genitalia similar to eu. Length 6-7.5 mm. Discussion: The arcuate shape of the lower lip of the female clypeal lobe (fig. 120), and the asymmetrically swollen flagellomeres of the male antenna (fig. 126) are diagnostic for euryops. The completely ridged propodeal hindface is an additional character that is shared only with lilIo. Females of euryops differ from eu and styphopteron in the essentially equal lower and upper interocular distances. The reduction of the second submarginal cell in the males from Argentina does not occur in the single female known from that country. Range: Pison euryops is known from Colombia, central and southern Brasil and northern Argentina. ~:
Holotype female: BRASIL, Santa Catarina: Nova Teutonia, XI-13-196I, 9, F. Piau mann (MCZ). Paratypes (24 specimens): COLOMBIA, Meta: Cord. Macarena, II-IS/29-76, 9, M. Cooper (BMNH). BRASIL, Distrito Federal: Res. Ecol. I8GE, various dates 1981-82, 4